Evolution
Overview
Evolution is any change across successive generation
s in the heritable
characteristics of biological
population
s. Evolutionary processes give rise to diversity at every level of biological organisation, including species
, individual organism
s and molecules
such as DNA
and protein
s.
Life on Earth originated
and then evolved from a universal common ancestor
approximately 3.7 billion years ago. Repeated speciation
and the divergence
of life can be traced through shared sets of biochemical and morphological traits, or by shared DNA sequences.
Generation
Generation , also known as procreation in biological sciences, is the act of producing offspring....
s in the heritable
Heritability
The Heritability of a population is the proportion of observable differences between individuals that is due to genetic differences. Factors including genetics, environment and random chance can all contribute to the variation between individuals in their observable characteristics...
characteristics of biological
Biology
Biology is a natural science concerned with the study of life and living organisms, including their structure, function, growth, origin, evolution, distribution, and taxonomy. Biology is a vast subject containing many subdivisions, topics, and disciplines...
population
Population
A population is all the organisms that both belong to the same group or species and live in the same geographical area. The area that is used to define a sexual population is such that inter-breeding is possible between any pair within the area and more probable than cross-breeding with individuals...
s. Evolutionary processes give rise to diversity at every level of biological organisation, including species
Biodiversity
Biodiversity is the degree of variation of life forms within a given ecosystem, biome, or an entire planet. Biodiversity is a measure of the health of ecosystems. Biodiversity is in part a function of climate. In terrestrial habitats, tropical regions are typically rich whereas polar regions...
, individual organism
Organism
In biology, an organism is any contiguous living system . In at least some form, all organisms are capable of response to stimuli, reproduction, growth and development, and maintenance of homoeostasis as a stable whole.An organism may either be unicellular or, as in the case of humans, comprise...
s and molecules
Molecular evolution
Molecular evolution is in part a process of evolution at the scale of DNA, RNA, and proteins. Molecular evolution emerged as a scientific field in the 1960s as researchers from molecular biology, evolutionary biology and population genetics sought to understand recent discoveries on the structure...
such as DNA
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...
and protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
s.
Life on Earth originated
Abiogenesis
Abiogenesis or biopoesis is the study of how biological life arises from inorganic matter through natural processes, and the method by which life on Earth arose...
and then evolved from a universal common ancestor
Last universal ancestor
The last universal ancestor , also called the last universal common ancestor , or the cenancestor, is the most recent organism from which all organisms now living on Earth descend. Thus it is the most recent common ancestor of all current life on Earth...
approximately 3.7 billion years ago. Repeated speciation
Speciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...
and the divergence
Anagenesis
Anagenesis, also known as "phyletic change," is the evolution of species involving an entire population rather than a branching event, as in cladogenesis. When enough mutations have occurred and become stable in a population so that it is significantly differentiated from an ancestral population,...
of life can be traced through shared sets of biochemical and morphological traits, or by shared DNA sequences.
Quotations
Anthropological, biological, and genetic evidence all put the origin of modern humans at between 200,000 and 100,000 years ago, probably in Africa. There is also much data that show an outburst of cultural behavior occurring around 50,000-40,000 years ago in Europe. That's when archaeologists date the oldest evidence of burial ceremonies, body ornaments, and cave paintings. 
William J. Cromie, Facing up to Modern Man; Harvard Gazette
I find it hard to swallow that I have only ten times more genes than those lowly bacteria in my gut. I had always liked the fact that they have ten thousand times less DNA than I did - that felt about right - but a factor of ten was carrying democracy a bit too far.
Gottfried Schatz in "Jeff's view on science and scientists", Amsterdam, Elsevier Butterworth-Heinemann, 2006, ISBN 978-0-444-52133-0, ISBN 0-444-52133-X (pbk.), p. 22, "Me and My Genome"
The most preposterous notion that H. sapiens has ever dreamed up is that the Lord God of Creation, Shaper and Ruler of all the Universes, wants the saccharine adoration of His creatures, can be swayed by their prayers, and becomes petulant if He does not receive this flattery. Yet this absurd fantasy, without a shred of evidence to bolster it, pays all the expenses of the oldest, largest, and least productive industry in all history.
Robert A. Heinlein, Time Enough for Love (1973)
[A] curious aspect of the theory of evolution is that everybody thinks he understands it.
Jacques Monod (1910-1979) On the Molecular Theory of Evolution (1974) (French Biochemist, Nobel Prize Medicine 1965)
Orgel's Second Rule: Evolution is cleverer than you are.
Francis Crick (British molecular biologist, 1916- ) quoted by Daniel C. Dennett in Elbow Room (1984)
"Evolution is a tinkerer.
Francois Jacob (French biochemist 1920- )"Evolution and Tinkering" (1977). See "Bricolage|Bricolage"
A hen is only an egg's way of making another egg.
Samuel Butler Life and Habit (1877)
Encyclopedia
Evolution is any change across successive generation
s in the heritable
characteristics of biological
population
s. Evolutionary processes give rise to diversity at every level of biological organisation, including species
, individual organism
s and molecules
such as DNA
and protein
s.
Life on Earth originated
and then evolved from a universal common ancestor
approximately 3.7 billion years ago. Repeated speciation
and the divergence
of life can be traced through shared sets of biochemical and morphological traits, or by shared DNA sequences. These homologous
traits and sequences are more similar among species that share a more recent common ancestor, and can be used to reconstruct
evolutionary histories, using both existing species and the fossil record. Existing patterns of biodiversity
have been shaped both by speciation
and by extinction
.
Charles Darwin
was the first to formulate a compelling scientific argument for the theory
of evolution by means of natural selection
. Evolution by natural selection is a process that is inferred
from three fact
s about populations: 1) more offspring are produced than can possibly survive, 2) traits vary among individuals, leading to differential rates of survival and reproduction, and 3) trait differences are heritable
. Thus, when members of a population die, they are replaced by individuals that are not born from random parents. Instead, these new members are born from parents that are better adapted
to the environment
in which natural selection took place. This can cause the evolution of traits that are seemingly fitted
for the functional
roles they perform. Natural selection is the only known cause of adaptation
, but not the only known cause of evolution. Other, nonadaptive causes of evolution
include mutation
and genetic drift
.
In the early 20th century, genetics
was integrated
with Darwin's theory of evolution by natural selection through the discipline of population genetics
. The importance of natural selection as a cause of evolution was accepted into other branches of biology
. Moreover, previously held notions about evolution, such as orthogenesis
and "progress"
became obsolete. Scientists continue to study evolution
by constructing theories
, by using observational data
, and by performing experiments in both the field
and the laboratory. Biologists agree
that descent with modification is one of the most reliably established facts in science. Discoveries in evolutionary biology have made a significant impact not just within the traditional branches of biology, but also in other academic disciplines (e.g., anthropology
and psychology
) and on society at large.
Greek philosophers, such as Anaximander and Empedocles. In contrast to these materialistic
views, Aristotle understood all natural things, not only living
things, as being imperfect actualization
s of different fixed natural possibilities, known as "forms
", "ideas
", or (in Latin translations) "species". This was part of his teleological
understanding of nature
in which all things have an intended role to play in a divine
cosmic
order. Variations of this idea became the standard understanding of the Middle Ages
, and were integrated into Christian learning, but Aristotle did not demand that real types of animals corresponded one-for-one with exact metaphysical forms, and specifically gave examples of how new types of living things could come to be.
In the 17th century the new method
of modern science rejected Aristotle's approach, and sought explanations of natural phenomena in terms of laws of nature which were the same for all visible things, and did not need to assume any fixed natural categories, nor any divine cosmic order. But this new approach was slow to take root in the biological sciences, which became the last bastion of the concept of fixed natural types. John Ray
used one of the previously more general terms for fixed natural types, "species", to apply to animal and plant types, but unlike Aristotle he strictly identified each type of living thing as a species, and proposed that each species can be defined by the features that perpetuate themselves each generation. These species were designed by God, but showing differences caused by local conditions. The biological classification introduced by Carolus Linnaeus
in 1735 also viewed species as fixed according to a divine plan.
Other naturalists of this time speculated on evolutionary change of species over time according to natural laws. Maupertuis
wrote in 1751 of natural modifications occurring during reproduction and accumulating over many generations to produce new species. Buffon
suggested that species could degenerate into different organisms, and Erasmus Darwin
proposed that all warm-blooded animals could have descended from a single micro-organism (or "filament"). The first fully-fledged evolutionary scheme was Lamarck's "transmutation" theory of 1809 which envisaged spontaneous generation continually producing simple forms of life developed greater complexity in parallel lineages with an inherent progressive tendency, and that on a local level these lineages adapted to the environment by inheriting changes caused by use or disuse in parents. (The latter process was later called Lamarckism.) These ideas were condemned by establishment naturalists as speculation lacking empirical support. In particular Georges Cuvier
insisted that species were unrelated and fixed, their similarities reflecting divine design for functional needs. In the meantime, Ray's ideas of benevolent design had been developed by William Paley
into a natural theology
which proposed complex adaptations as evidence of divine design, and was admired by Charles Darwin.
The critical break from the concept of fixed species in biology began with the theory of evolution by natural selection, which was formulated by Charles Darwin. Partly influenced by An Essay on the Principle of Population
by Thomas Robert Malthus, Darwin noted that population growth would lead to a "struggle for existence" where favorable variations could prevail as others perished. Each generation, many offspring fail to survive to an age of reproduction because of limited resources. This could explain the diversity of animals and plants from a common ancestry through the working of natural laws working the same for all types of thing. Darwin was developing his theory of "natural selection
" from 1838 onwards until Alfred Russel Wallace
sent him a similar theory in 1858. Both men presented their separate papers to the Linnean Society of London
. At the end of 1859, Darwin's publication of On the Origin of Species explained natural selection in detail and in a way that lead to an increasingly wide acceptance of Darwinian evolution
. Thomas Henry Huxley applied Darwin's ideas to humans, using paleontology
and comparative anatomy
to provide strong evidence that humans and apes shared a common ancestry. Some were disturbed by this since it implied that humans did not have a special place in the universe.
Precise mechanisms of reproductive heritability and the origin of new traits remained a mystery. Towards this end, Darwin developed his provisional theory of pangenesis
. In 1865 Gregor Mendel
reported that traits were inherited in a predictable manner through the independent assortment and segregation of elements (later known as genes
). Mendel's laws of inheritance eventually supplanted most of Darwin's pangenesis theory. August Weismann
made the important distinction between germ cells (sperm and eggs)
and somatic cells of the body, demonstrating that heredity passes through the germ line only. Hugo de Vries
connected Darwin's pangenesis theory to Wiesman's germ/soma cell distinction and proposed that Darwin's pangenes were concentrated in the cell nucleus
and when expressed they could move into the cytoplasm
to change the cells structure. De Vries was also one of the researchers who made Mendel's work well-known, believing that Mendelian traits corresponded to the transfer of heritable variations along the germline. To explain how new variants originate, De Vries developed a mutation
theory that led to a temporary rift between those who accepted Darwinian evolution and biometricians who allied with de Vries. At the turn of the 20th century, pioneers in the field of population genetics, such as J.B.S. Haldane, Sewall Wright
, and Ronald Fisher
, set the foundations of evolution onto a robust statistical philosophy. The false contradiction between Darwin's theory, genetic mutations, and Mendelian inheritance was thus reconciled.
In the 1920s and 1930s a modern evolutionary synthesis connected natural selection, mutation theory, and Mendelian inheritance into a unified theory that applied generally to any branch of biology. The modern synthesis was able to explain patterns observed across species in populations, through fossil transitions in palaeontology, and even complex cellular mechanisms in developmental biology
. The publication of the structure of DNA by James Watson
and Francis Crick
in 1953 demonstrated a physical basis for inheritance. Molecular biology
improved our understanding of the relationship between genotype and phenotype. 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
. In 1973, evolutionary biologist Theodosius Dobzhansky
penned that "nothing in biology makes sense except in the light of evolution", because it has brought to light the relations of what first seemed disjointed facts in natural history into a coherent explanatory body of knowledge that describes and predicts many observable facts about life on this planet.
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. This extension has been dubbed "eco-evo-devo
".
– particular 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. Inherited traits are controlled by gene
s and the complete set of genes within an organism's genome
is called its genotype
.
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
. As a result, many aspects of an organism's phenotype are not inherited. 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 their genotype; 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 known to be passed from one generation to the next via DNA
, a molecule
that encodes genetic information. DNA is a long polymer
composed of four types of bases. 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. Before a cell divides, the DNA is copied, so that each of the resulting two cells will inherit the DNA sequence. Portions of a DNA molecule that specify a single functional unit are called gene
s; different genes have different sequences of bases. Within cells
, the long strands of DNA form condensed structures called chromosome
s. 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 allele
s. DNA sequences can change through mutation
s, 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. However, while this simple correspondence between an allele and a trait works in some cases, most traits are more complex and are controlled by multiple interacting genes
.
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. DNA methylation
marking chromatin
, self-sustaining metabolic loops, gene silencing by RNA interference
and the three dimensional conformation
of proteins (such as prions) are areas where epigenetic inheritance systems have been discovered at the organismic level. Developmental biologists suggest that complex interactions in genetic networks
and communication among cells can lead to heritable variations that may underlay some of the mechanics in developmental plasticity
and canalization
. Heritability may also occur at even larger scales. 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. Other examples of heritability in evolution that are not under the direct control of genes include the inheritance of cultural traits
and symbiogenesis
.
results from both its genotype
and the influence from the environment
it has lived in. A substantial part of the variation in phenotypes in a population is caused by the differences between their genotypes. The modern evolutionary synthesis
defines evolution as the change over time in this genetic 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 mutation
s in genetic material, 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. However, even relatively small differences in genotype can lead to dramatic differences in phenotype: for example, chimpanzees and humans differ in only about 5% of their genomes.
, or prevent the gene from functioning. Based on studies in the fly Drosophila melanogaster
, it has been suggested that if a mutation changes a protein produced by a gene, this will probably be harmful, with about 70% of these mutations having damaging effects, and the remainder being either neutral or weakly beneficial.
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. Extra copies of genes are a major source of the raw material needed for new genes to evolve. This is important because most new genes evolve within gene families
from pre-existing genes that share common ancestors. For example, the human eye uses four genes to make structures that sense light: three for colour vision
and one for night vision
; all four are descended from a single ancestral gene.
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. Other types of mutations can even generate entirely new genes from previously noncoding DNA.
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. When new genes are assembled from shuffling pre-existing parts, domains
act as modules with simple independent functions, which can be mixed together to produce new combinations with new and complex functions. For example, polyketide synthase
s are large enzymes that make antibiotics; they contain up to one hundred independent domains that each catalyze one step in the overall process, like a step in an assembly line.
ual 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. Recombination and reassortment do not alter allele frequencies, but instead change which alleles are associated with each other, producing offspring with new combinations of alleles. Sex usually increases genetic variation and may increase the rate of evolution.
is the exchange of genes between populations and between species. It can therefore be a source of variation that is new to a population or to a 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
. In medicine, this contributes to the spread of antibiotic resistance
, as when one bacteria acquires resistance genes it can rapidly transfer them to other species. Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces cerevisiae
and the adzuki bean beetle Callosobruchus chinensis has occurred. An example of larger-scale transfers are the eukaryotic bdelloid rotifers, which have received a range of genes from bacteria, fungi and plants. Virus
es can also carry DNA between organisms, allowing transfer of genes even across biological domains
.
Large-scale gene transfer has also occurred between the ancestors of eukaryotic cells
and bacteria
, during the acquisition of chloroplast
s and mitochondria
. It is possible that eukaryotes themselves originated from horizontal gene transfers between bacteria
and archaea
.
, genetic drift
, genetic hitchhiking
, mutation
and gene flow
.
is the process by which genetic mutations that enhance reproduction become and remain, 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:
These conditions produce competition between organisms for survival and reproduction. Consequently, organisms with traits that give them an advantage over their competitors pass these advantageous traits on, while traits that do not confer an advantage are not passed on to the next generation.
The central concept of natural selection is the evolutionary fitness
of an organism. Fitness is measured by an organism's ability to survive and reproduce, which determines the size of its genetic contribution to the next generation. However, fitness is not the same as the total number of offspring: instead fitness is indicated by the proportion of subsequent generations that carry an organism's genes. For example, if an organism could survive well and reproduce rapidly, but its offspring were all too small and weak to survive, this organism would make little genetic contribution to future generations and would thus have low fitness.
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". Examples of traits that can increase fitness are enhanced survival and increased fecundity
. Conversely, the lower fitness caused by having a less beneficial or deleterious allele results in this allele becoming rarer — they are "selected against". Importantly, the fitness of an allele is not a fixed characteristic; if the environment changes, previously neutral or harmful traits may become beneficial and previously beneficial traits become harmful. However, even if the direction of selection does reverse in this way, traits that were lost in the past may not re-evolve in an identical form (see Dollo's law
).
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. Secondly, disruptive selection
is selection for extreme trait values and often results in two different values
becoming most common, with selection against the average value. This would be when either short or tall organisms had an advantage, but not those of medium height. Finally, in stabilizing selection
there is selection against extreme trait values on both ends, which causes a decrease in variance
around the average value and less diversity. This would, for example, cause organisms to slowly become all the same height.
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. Traits that evolved through sexual selection are particularly prominent in males of some animal species, despite traits such as cumbersome antlers, mating calls or bright colours that attract predators, decreasing the survival of individual males. This survival disadvantage is balanced by higher reproductive success in males that show these hard to fake
, sexually selected traits.
Natural selection most generally makes nature the measure against which individuals and individual traits, are more or less likely to survive. "Nature" in this sense refers to an ecosystem
, that is, a system in which organisms interact with every other element, physical as well as biological
, in their local environment
. Eugene Odum, a founder of ecology, defined an ecosystem as: "Any unit that includes all of the organisms...in a given area interacting with the physical environment so that a flow of energy leads to clearly defined trophic structure, biotic diversity and material cycles (ie: exchange of materials between living and nonliving parts) within the system." Each population within an ecosystem occupies a distinct niche
, or position, with distinct relationships to other parts of the system. 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. Selection can act at multiple levels simultaneously. An example of selection occurring below the level of the individual organism are genes called transposon
s, which can replicate and spread throughout a genome
. Selection at a level above the individual, such as group selection
, may allow the evolution of co-operation, as discussed below.
evolution. For example, according to the phenotype-first theory of evolution
, mutations can eventually cause the genetic assimilation
of traits that were previously induced by the environment
.
Mutation bias effects are superimposed on other processes. If selection would favor 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. Mutations leading to the loss of function of a gene are much more common than mutations that produce a new, fully functional gene. Most loss of function mutations are selected against. But when selection is weak, mutation bias towards loss of function can affect evolution. For example, pigment
s are no longer useful when animals live in the darkness of caves, and tend to be lost. This kind of loss of function can occur because of mutation bias, and/or because the function had a cost, and once the benefit of the function disappeared, natural selection leads to the loss. Loss of sporulation
ability in a bacterium
during laboratory evolution appears to have been caused by mutation bias, rather than natural selection against the cost of maintaining sporulation ability. When there is no selection for loss of function, the speed at which loss evolves depends more on the mutation rate than it does on the effective population size
, indicating that it is driven more by mutation bias than by genetic drift.
from one generation to the next that occurs because alleles are subject to sampling error
. As a result, when selective forces are absent or relatively weak, allele frequencies tend to "drift" upward or downward randomly (in a random walk
). This drift halts when an allele eventually becomes fixed
, either by disappearing from the population, or replacing the other alleles entirely. 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.
It is usually difficult to measure the relative importance of selection and neutral processes, including drift. The comparative importance of adaptive and non-adaptive forces in driving evolutionary change is an area of current research
.
The neutral theory of molecular evolution
proposed that most evolutionary changes are the result of the fixation of neutral mutation
s by genetic drift. Hence, in this model, most genetic changes in a population are the result of constant mutation pressure and genetic drift. This form of the neutral theory is now largely abandoned, since it does not seem to fit the genetic variation seen in nature. However, a more recent and better-supported version of this model is the nearly neutral theory
, where a mutation that would be neutral in a small population is not necessarily neutral in a large population. Other alternative theories propose that genetic drift is dwarfed by other stochastic forces in evolution, such as genetic hitchhiking
, also known as genetic draft.
The time for a neutral allele to become fixed by genetic drift depends on population size, with fixation occurring more rapidly in smaller populations. The number of individuals in a population is not critical, but instead a measure known as the effective population size
. The effective population is usually smaller than the total population since it takes into account factors such as the level of inbreeding and the stage of the lifecycle in which the population is the smallest. The effective population size may not be the same for every gene in the same population.
. This tendency is measured by finding how often two alleles occur together on a single chromosome compared to expectations, which is called their linkage disequilibrium
. 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: natural selection can drive a selective sweep
that will also cause the other alleles in the haplotype to become more common in the population; this effect is called genetic hitchhiking
or genetic draft. Genetic draft caused by the fact that some neutral genes are genetically linked to others that are under selection can be partially captured by an appropriate effective population size.
is the exchange of genes between populations and between species. The presence or absence of gene flow fundamentally changes the course of evolution. 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 organism within these populations to evolve 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
's developed his shifting balance theory
that gene flow between partially isolated populations was an important aspect of adaptive evolution. However, recently there has been substantial criticism of the importance of the shifting balance theory
.
s 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 co-operating
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 sometimes divided into macroevolution
, which is evolution that occurs at or above the level of species, such as extinction
and speciation
and microevolution
, which is smaller evolutionary changes, such as adaptations, within a species or population. In general, macroevolution is regarded as the outcome of long periods of microevolution. Thus, the distinction between micro- and macroevolution is not a fundamental one – the difference is simply the time involved. However, in macroevolution, the traits of the entire species may be important. 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 or long-term plans; realistically however, evolution has no long-term goal and does not necessarily produce greater complexity. Although complex species
have evolved, they occur as a side effect of the overall number of organisms increasing and simple forms of life still remain more common in the biosphere. For example, the overwhelming majority of species are microscopic prokaryote
s, which form about half the world's biomass
despite their small size, and constitute the vast majority of Earth's biodiversity. Simple organisms have therefore been the dominant form of life on Earth throughout its history and continue to be the main form of life up to the present day, with complex life only appearing more diverse because it is more noticeable
. Indeed, the evolution of microorganism
s is particularly important to modern evolutionary research
, since their rapid reproduction allows the study of experimental evolution
and the observation of evolution and adaptation in real time.
. Also, the term adaptation may refer to a trait
that is important for an organism's survival. 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
. The following definitions are due to Theodosius Dobzhansky
.
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. Other striking examples are the bacteria Escherichia coli
evolving the ability to use citric acid
as a nutrient in a long-term laboratory experiment
, Flavobacterium
evolving a novel enzyme that allows these bacteria to grow on the by-products of nylon
manufacturing, and the soil bacterium Sphingobium
evolving an entirely new metabolic pathway
that degrades the synthetic pesticide
pentachlorophenol
. An interesting but still controversial idea is that some adaptations might increase the ability of organisms to generate genetic diversity and adapt by natural selection (increasing organisms' evolvability
).
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. However, since all living organisms are related to some extent, even organs that appear to have little or no structural similarity, such as arthropod
, squid and vertebrate eyes, or the limbs and wings of arthropods and vertebrates, can depend on a common set of homologous genes that control their assembly and function; this is called deep homology
.
During evolution, some structures may lose their original function and become vestigial structure
s. Such structures may have little or no function in a current species, yet have a clear function in ancestral species, or other closely related species. Examples include pseudogene
s, the non-functional remains of eyes in blind cave-dwelling fish, wings in flightless birds, and the presence of hip bones in whales and snakes. Examples of vestigial structures in humans
include wisdom teeth, the coccyx
, the vermiform appendix
, and other behavioural vestiges such as goose bumps
and primitive reflexes
.
However, many traits that appear to be simple adaptations are in fact exaptation
s: structures originally adapted for one function, but which coincidentally became somewhat useful for some other function in the process. One example is the African lizard Holaspis guentheri, which developed an extremely flat head for hiding in crevices, as can be seen by looking at its near relatives. However, in this species, the head has become so flattened that it assists in gliding from tree to tree—an exaptation
. Within cells, molecular machine
s such as the bacterial flagella and protein sorting machinery
evolved by the recruitment of several pre-existing proteins that previously had different functions. Another example is the recruitment of enzymes from glycolysis
and xenobiotic metabolism
to serve as structural proteins called crystallin
s within the lenses of organisms' eye
s.
A critical principle of ecology
is that of competitive exclusion
: no two species can occupy the same niche in the same environment for a long time. Consequently, natural selection will tend to force species to adapt to different ecological niche
s. This may mean that, for example, two species of cichlid
fish adapt to live in different habitat
s, which will minimise the competition between them for food.
An area of current investigation in evolutionary developmental biology
is the developmental
basis of adaptations and exaptations. This research addresses the origin and evolution of embryonic development
and how modifications of development and developmental processes produce novel features. These studies have shown that evolution can alter development to produce new structures, such as embryonic bone structures that develop into the jaw in other animals instead forming part of the middle ear in mammals. It is also possible for structures that have been lost in evolution to reappear due to changes in developmental genes, such as a mutation in chicken
s causing embryos to grow teeth similar to those of crocodile
s. It is now becoming clear that most alterations in the form of organisms are due to changes in a small set of conserved genes.
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 co-evolution
. An example is the production of tetrodotoxin
in the rough-skinned newt
and the evolution of tetrodotoxin resistance in its predator, the common garter snake
. 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.
that grow on their roots and aid the plant in absorbing nutrients from the soil. This is a reciprocal
relationship as the plants provide the fungi with sugars from photosynthesis. 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 bee
s, termite
s and ant
s, 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 cell
s 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 cell
s 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. This activity is selected for because if the helping individual contains alleles which promote the helping activity, it is likely that its kin will also contain these alleles and thus those alleles will be passed on. Other processes that may promote cooperation include group selection
, where cooperation provides benefits to a group of organisms.
is the process where a species diverges into two or more descendant species.
There are multiple ways to define the concept of "species". The choice of definition is dependent on the particularities of the species concerned. For example, some species concepts apply more readily toward sexually reproducing organisms while others lend themselves better toward asexual organisms. Despite the diversity of various species concepts, these various concepts can be placed into one of three broad philosophical approaches: interbreeding, ecological and phylogenetic. The biological species concept (BSC) is a classic example of the interbreeding approach. Defined by Ernst Mayr in 1942, the BSC states that "species are groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups". Despite its wide and long-term use, the BSC like others is not without controversy, for example because these concepts cannot be applied to prokaryotes, and this is called the species problem
. Some researchers have attempted a unifying monistic definition of species, while others adopt a pluralistic approach and suggest that there may be a different ways to logically interpret the definition of a species. "
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 horse
s and donkey
s mating to produce mule
s. Such hybrids are generally infertile
. 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. The importance of hybridisation in producing new species
of animals is unclear, although cases have been seen in many types of animals, with the gray tree frog being a particularly well-studied example.
Speciation has been observed multiple times under both controlled laboratory conditions and in nature. In sexually reproducing organisms, speciation results from reproductive isolation followed by genealogical divergence. There are four mechanisms for 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. As selection and drift act independently on populations isolated from the rest of their species, separation may eventually produce organisms that cannot interbreed.
The second mechanism 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 mechanism of speciation 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. Generally this occurs when there has been a drastic change in the environment within the parental species' habitat. One example is the grass Anthoxanthum odoratum
, which can undergo parapatric speciation in response to localised metal pollution from mines. Here, plants evolve that have resistance to high levels of metals in the soil. 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. Generally, sympatric speciation in animals requires the evolution of both genetic differences
and non-random mating
, to allow reproductive isolation to evolve.
One type of sympatric speciation involves cross-breeding 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 chromosome
s 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
. This allows the chromosomes from each parental species to form matching pairs during meiosis, since each parent's chromosomes are represented by a pair already. An example of such a speciation event is when the plant species Arabidopsis thaliana
and Arabidopsis arenosa cross-bred to give the new species Arabidopsis suecica. This happened about 20,000 years ago, and the speciation process has been repeated in the laboratory, which allows the study of the genetic mechanisms involved in this process. Indeed, chromosome doubling within a species may be a common cause of reproductive isolation, as half the doubled chromosomes will be unmatched when breeding with undoubled organisms.
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. In this theory, speciation and rapid evolution are linked, with natural selection and genetic drift acting most strongly on organisms undergoing speciation in novel habitats or small populations. 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. Nearly all animal and plant species that have lived on Earth are now extinct, and extinction appears to be the ultimate fate of all species. These extinctions have happened continuously throughout the history of life, although the rate of extinction spikes in occasional mass extinction event
s. The Cretaceous–Tertiary extinction event, during which the non-avian dinosaurs went extinct, is the most well-known, but the earlier Permian–Triassic extinction event was even more severe, with approximately 96% of species driven to extinction. The Holocene extinction event
is an ongoing mass extinction associated with humanity's expansion across the globe over the past few thousand years. Present-day extinction rates are 100–1000 times greater than the background rate and up to 30% of species may be extinct by the mid 21st century. Human activities are now the primary cause of the ongoing extinction event; global warming
may further accelerate it in the future.
The role of extinction in evolution is not very well understood and may depend on which type of extinction is considered. The causes of the continuous "low-level" extinction events, which form the majority of extinctions, may be the result of competition between species for limited resources (competitive exclusion). If one species can out-compete another, this could produce species selection, with the fitter species surviving and the other species being driven to extinction. The intermittent mass extinctions are also important, but instead of acting as a selective force, they drastically reduce diversity in a nonspecific manner and promote bursts of rapid evolution
and speciation in survivors.
is that the complex biochemistry
that makes up life came from simpler chemical reactions. The beginning of life may have included self-replicating molecules such as RNA
, and the assembly of simple cells.
s on Earth
are descended from a common ancestor or ancestral gene pool. Current species are a stage in the process of evolution, with their diversity the product of a long series of speciation and extinction events. The common descent
of organisms was first deduced from four simple facts about organisms: 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. However, modern research has suggested that, due to horizontal gene transfer, this "tree of life
" may be more complicated than a simple branching tree since some genes have spread independently between distantly related species.
Past species have also left records of their evolutionary history. Fossil
s, along with the comparative anatomy of present-day organisms, constitute the morphological, or anatomical, record. By comparing the anatomies of both modern and extinct species, paleontologists can infer the lineages of those species. 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 nucleotide
s and amino acid
s. The development of molecular genetics
has revealed the record of evolution left in organisms' genome
s: dating when species diverged through the molecular clock
produced by mutations. For example, these DNA sequence comparisons have revealed that humans and chimpanzees share 96% of their genomes and analyzing the few areas where they differ helps shed light on when the common ancestor of these species existed.
s inhabited the Earth from approximately 3–4 billion years ago. No obvious changes in morphology
or cellular organisation occurred in these organisms over the next few billion years.
The eukaryotic cells
emerged between 1.6 – 2.7 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 engulfed bacteria and the host cell then underwent co-evolution, with the bacteria evolving into either mitochondria
or hydrogenosome
s. Another engulfment of cyanobacterial-like organisms led to the formation of chloroplast
s in algae and plants.
The history of life was that of the unicellular eukaryotes, prokaryotes and archaea until about 610 million years ago when multicellular organisms began to appear in the oceans in the Ediacaran
period. The evolution of multicellularity occurred in multiple independent events, in organisms as diverse as sponges, brown algae
, cyanobacteria, slime moulds and myxobacteria
.
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. Various triggers for the Cambrian explosion have been proposed, including the accumulation of oxygen
in the atmosphere
from photosynthesis
.
About 500 million years ago, plant
s and fungi
colonised the land and were soon followed by arthropod
s and other animals. Insect
s were particularly successful and even today make up the majority of animal species. Amphibian
s first appeared around 364 million years ago, followed by early amniote
s, then bird
s around 155 million years ago (both from "reptile
"-like lineages), mammal
s around 129 million years ago, homininae
around 10 million years ago and modern humans around 0.25 million years ago. 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.
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. More recently, such selection has become a vital part of genetic engineering
, with selectable marker
s such as antibiotic resistance genes being used to manipulate DNA. In repeated rounds of mutation and selection proteins with valuable properties have evolved, for example modified enzyme
s and new antibodies
, in a process called directed evolution
.
Understanding the changes that have occurred during organism's evolution can reveal the genes needed to construct parts of the body, genes which may be involved in human genetic disorder
s. For example, the mexican tetra
is an albino cavefish that lost its eyesight during evolution. 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. This helped identify genes required for vision and pigmentation.
In computer science
, simulations of evolution using evolutionary algorithm
s and artificial life
started in the 1960s and was extended with simulation of artificial selection
. Artificial evolution
became a widely recognised optimisation method as a result of the work of Ingo Rechenberg
in the 1960s. He used evolution strategies
to solve complex engineering problems. Genetic algorithm
s in particular became popular through the writing of John Holland
. Practical applications also include automatic evolution of computer programs
. Evolutionary algorithms are now used to solve multi-dimensional problems more efficiently than software produced by human designers and also to optimise the design of systems.
.
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 religion
s and who raise various objections to evolution
. As had been demonstrated by responses to the publication of Vestiges of the Natural History of Creation
in 1844, the most controversial aspect of evolutionary biology is the implication of human evolution
that humans share common ancestry with apes and that the mental and moral faculties of humanity have the same types of natural causes as other inherited traits in animals. In some countries, notably the United States, these tensions between science and religion have fuelled the current creation-evolution controversy, a religious conflict focusing on politics
and public education
. While other scientific fields such as cosmology
and Earth science
also conflict with literal interpretations of many religious texts, evolutionary biology experiences significantly more opposition from religious literalists.
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 1925 caused the subject to become very rare in American secondary biology textbooks for a generation, but it was gradually re-introduced about a generation later and legally protected with the 1968 Epperson v. Arkansas
decision. Since then, the competing religious belief of creationism was legally disallowed in secondary school curricula in various decisions in the 1970s and 1980s, but it returned in the form of intelligent design
, to be excluded once again in the 2005 Kitzmiller v. Dover Area School District
case.
Generation
Generation , also known as procreation in biological sciences, is the act of producing offspring....
s in the heritable
Heritability
The Heritability of a population is the proportion of observable differences between individuals that is due to genetic differences. Factors including genetics, environment and random chance can all contribute to the variation between individuals in their observable characteristics...
characteristics of biological
Biology
Biology is a natural science concerned with the study of life and living organisms, including their structure, function, growth, origin, evolution, distribution, and taxonomy. Biology is a vast subject containing many subdivisions, topics, and disciplines...
population
Population
A population is all the organisms that both belong to the same group or species and live in the same geographical area. The area that is used to define a sexual population is such that inter-breeding is possible between any pair within the area and more probable than cross-breeding with individuals...
s. Evolutionary processes give rise to diversity at every level of biological organisation, including species
Biodiversity
Biodiversity is the degree of variation of life forms within a given ecosystem, biome, or an entire planet. Biodiversity is a measure of the health of ecosystems. Biodiversity is in part a function of climate. In terrestrial habitats, tropical regions are typically rich whereas polar regions...
, individual organism
Organism
In biology, an organism is any contiguous living system . In at least some form, all organisms are capable of response to stimuli, reproduction, growth and development, and maintenance of homoeostasis as a stable whole.An organism may either be unicellular or, as in the case of humans, comprise...
s and molecules
Molecular evolution
Molecular evolution is in part a process of evolution at the scale of DNA, RNA, and proteins. Molecular evolution emerged as a scientific field in the 1960s as researchers from molecular biology, evolutionary biology and population genetics sought to understand recent discoveries on the structure...
such as DNA
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...
and protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
s.
Life on Earth originated
Abiogenesis
Abiogenesis or biopoesis is the study of how biological life arises from inorganic matter through natural processes, and the method by which life on Earth arose...
and then evolved from a universal common ancestor
Last universal ancestor
The last universal ancestor , also called the last universal common ancestor , or the cenancestor, is the most recent organism from which all organisms now living on Earth descend. Thus it is the most recent common ancestor of all current life on Earth...
approximately 3.7 billion years ago. Repeated speciation
Speciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...
and the divergence
Anagenesis
Anagenesis, also known as "phyletic change," is the evolution of species involving an entire population rather than a branching event, as in cladogenesis. When enough mutations have occurred and become stable in a population so that it is significantly differentiated from an ancestral population,...
of life can be traced through shared sets of biochemical and morphological traits, or by shared DNA sequences. These homologous
Homology (biology)
Homology forms the basis of organization for comparative biology. In 1843, Richard Owen defined homology as "the same organ in different animals under every variety of form and function". Organs as different as a bat's wing, a seal's flipper, a cat's paw and a human hand have a common underlying...
traits and sequences are more similar among species that share a more recent common ancestor, and can be used to reconstruct
Phylogenetics
In biology, phylogenetics is the study of evolutionary relatedness among groups of organisms , which is discovered through molecular sequencing data and morphological data matrices...
evolutionary histories, using both existing species and the fossil record. Existing patterns of biodiversity
Biodiversity
Biodiversity is the degree of variation of life forms within a given ecosystem, biome, or an entire planet. Biodiversity is a measure of the health of ecosystems. Biodiversity is in part a function of climate. In terrestrial habitats, tropical regions are typically rich whereas polar regions...
have been shaped both by speciation
Speciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...
and by extinction
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...
.
Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...
was the first to formulate a compelling scientific argument for the theory
Scientific theory
A scientific theory comprises a collection of concepts, including abstractions of observable phenomena expressed as quantifiable properties, together with rules that express relationships between observations of such concepts...
of evolution by means of natural selection
Natural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....
. Evolution by natural selection is a process that is inferred
Inference
Inference is the act or process of deriving logical conclusions from premises known or assumed to be true. The conclusion drawn is also called an idiomatic. The laws of valid inference are studied in the field of logic.Human inference Inference is the act or process of deriving logical conclusions...
from three fact
Fact
A fact is something that has really occurred or is actually the case. The usual test for a statement of fact is verifiability, that is whether it can be shown to correspond to experience. Standard reference works are often used to check facts...
s about populations: 1) more offspring are produced than can possibly survive, 2) traits vary among individuals, leading to differential rates of survival and reproduction, and 3) trait differences are heritable
Heritability
The Heritability of a population is the proportion of observable differences between individuals that is due to genetic differences. Factors including genetics, environment and random chance can all contribute to the variation between individuals in their observable characteristics...
. Thus, when members of a population die, they are replaced by individuals that are not born from random parents. Instead, these new members are born from parents that are better adapted
Adaptation
An adaptation in biology is a trait with a current functional role in the life history of an organism that is maintained and evolved by means of natural selection. An adaptation refers to both the current state of being adapted and to the dynamic evolutionary process that leads to the adaptation....
to the environment
Environment (biophysical)
The biophysical environment is the combined modeling of the physical environment and the biological life forms within the environment, and includes all variables, parameters as well as conditions and modes inside the Earth's biosphere. The biophysical environment can be divided into two categories:...
in which natural selection took place. This can cause the evolution of traits that are seemingly fitted
Teleonomy
Teleonomy is the quality of apparent purposefulness and of goal-directedness of structures and functions in living organisms that derive from their evolutionary history, adaptation for reproductive success, or generally, due to the operation of a program....
for the functional
Function (biology)
A function is part of an answer to a question about why some object or process occurred in a system that evolved through a process of selection. Thus, function refers forward from the object or process, along some chain of causation, to the goal or success...
roles they perform. Natural selection is the only known cause of adaptation
Adaptation
An adaptation in biology is a trait with a current functional role in the life history of an organism that is maintained and evolved by means of natural selection. An adaptation refers to both the current state of being adapted and to the dynamic evolutionary process that leads to the adaptation....
, but not the only known cause of evolution. Other, nonadaptive causes of evolution
Microevolution
Microevolution is the changes in allele frequencies that occur over time within a population. This change is due to four different processes: mutation, selection , gene flow, and genetic drift....
include mutation
Mutation
In molecular biology and genetics, mutations are changes in a genomic sequence: the DNA sequence of a cell's genome or the DNA or RNA sequence of a virus. They can be defined as sudden and spontaneous changes in the cell. Mutations are caused by radiation, viruses, transposons and mutagenic...
and genetic drift
Genetic drift
Genetic drift or allelic drift is the change in the frequency of a gene variant in a population due to random sampling.The alleles in the offspring are a sample of those in the parents, and chance has a role in determining whether a given individual survives and reproduces...
.
In the early 20th century, genetics
Classical genetics
Classical genetics consists of the technique and methodologies of genetics that predate the advent of molecular biology. A key discovery of classical genetics in eukaryotes was genetic linkage...
was integrated
Modern evolutionary synthesis
The modern evolutionary synthesis is a union of ideas from several biological specialties which provides a widely accepted account of evolution...
with Darwin's theory of evolution by natural selection through the discipline of population genetics
Population genetics
Population genetics is the study of allele frequency distribution and change under the influence of the four main evolutionary processes: natural selection, genetic drift, mutation and gene flow. It also takes into account the factors of recombination, population subdivision and population...
. The importance of natural selection as a cause of evolution was accepted into other branches of biology
Biology
Biology is a natural science concerned with the study of life and living organisms, including their structure, function, growth, origin, evolution, distribution, and taxonomy. Biology is a vast subject containing many subdivisions, topics, and disciplines...
. Moreover, previously held notions about evolution, such as orthogenesis
Orthogenesis
Orthogenesis, orthogenetic evolution, progressive evolution or autogenesis, is the hypothesis that life has an innate tendency to evolve in a unilinear fashion due to some internal or external "driving force". The hypothesis is based on essentialism and cosmic teleology and proposes an intrinsic...
and "progress"
Largest-scale trends in evolution
The history of life on Earth seems to show a clear trend; for example, it seems intuitive that there is a trend towards increasing complexity in living organisms. More recent organisms, such as mammals, appear to be much more complex than older organisms, such as bacteria. However, there are...
became obsolete. Scientists continue to study evolution
Current research in evolutionary biology
Current research in evolutionary biology covers diverse topics, as should be expected given the centrality of evolution to understanding biology...
by constructing theories
Mathematical and theoretical biology
Mathematical and theoretical biology is an interdisciplinary scientific research field with a range of applications in biology, medicine and biotechnology...
, by using observational data
Observational science
An observational science is a science where it is not possible to construct controlled experiments in the area under study. For example, in astronomy, it is not possible to create or manipulate stars or galaxies in order to observe what happens...
, and by performing experiments in both the field
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...
and the laboratory. Biologists agree
Scientific consensus
Scientific consensus is the collective judgment, position, and opinion of the community of scientists in a particular field of study. Consensus implies general agreement, though not necessarily unanimity. Scientific consensus is not by itself a scientific argument, and it is not part of the...
that descent with modification is one of the most reliably established facts in science. Discoveries in evolutionary biology have made a significant impact not just within the traditional branches of biology, but also in other academic disciplines (e.g., anthropology
Biological anthropology
Biological anthropology is that branch of anthropology that studies the physical development of the human species. It plays an important part in paleoanthropology and in forensic anthropology...
and psychology
Evolutionary psychology
Evolutionary psychology is an approach in the social and natural sciences that examines psychological traits such as memory, perception, and language from a modern evolutionary perspective. It seeks to identify which human psychological traits are evolved adaptations, that is, the functional...
) and on society at large.
History of evolutionary thought
The proposal that one type of animal could descend from an animal of another type goes back to some of the first pre-SocraticPre-Socratic philosophy
Pre-Socratic philosophy is Greek philosophy before Socrates . In Classical antiquity, the Presocratic philosophers were called physiologoi...
Greek philosophers, such as Anaximander and Empedocles. In contrast to these materialistic
Materialism
In philosophy, the theory of materialism holds that the only thing that exists is matter; that all things are composed of material and all phenomena are the result of material interactions. In other words, matter is the only substance...
views, Aristotle understood all natural things, not only living
Life
Life is a characteristic that distinguishes objects that have signaling and self-sustaining processes from those that do not, either because such functions have ceased , or else because they lack such functions and are classified as inanimate...
things, as being imperfect actualization
Actuality
Actuality may refer to:* Potentiality and actuality * Actuality film...
s of different fixed natural possibilities, known as "forms
Theory of Forms
Plato's theory of Forms or theory of Ideas asserts that non-material abstract forms , and not the material world of change known to us through sensation, possess the highest and most fundamental kind of reality. When used in this sense, the word form is often capitalized...
", "ideas
Idealism
In philosophy, idealism is the family of views which assert that reality, or reality as we can know it, is fundamentally mental, mentally constructed, or otherwise immaterial. Epistemologically, idealism manifests as a skepticism about the possibility of knowing any mind-independent thing...
", or (in Latin translations) "species". This was part of his teleological
Teleology
A teleology is any philosophical account which holds that final causes exist in nature, meaning that design and purpose analogous to that found in human actions are inherent also in the rest of nature. The word comes from the Greek τέλος, telos; root: τελε-, "end, purpose...
understanding of nature
Nature (philosophy)
Nature is a concept with two major sets of inter-related meanings, referring on the one hand to the things which are natural, or subject to the normal working of "laws of nature", or on the other hand to the essential properties and causes of those things to be what they naturally are, or in other...
in which all things have an intended role to play in a divine
Divinity
Divinity and divine are broadly applied but loosely defined terms, used variously within different faiths and belief systems — and even by different individuals within a given faith — to refer to some transcendent or transcendental power or deity, or its attributes or manifestations in...
cosmic
Cosmos
In the general sense, a cosmos is an orderly or harmonious system. It originates from the Greek term κόσμος , meaning "order" or "ornament" and is antithetical to the concept of chaos. Today, the word is generally used as a synonym of the word Universe . The word cosmos originates from the same root...
order. Variations of this idea became the standard understanding of the Middle Ages
Middle Ages
The Middle Ages is a periodization of European history from the 5th century to the 15th century. The Middle Ages follows the fall of the Western Roman Empire in 476 and precedes the Early Modern Era. It is the middle period of a three-period division of Western history: Classic, Medieval and Modern...
, and were integrated into Christian learning, but Aristotle did not demand that real types of animals corresponded one-for-one with exact metaphysical forms, and specifically gave examples of how new types of living things could come to be.
In the 17th century the new method
Scientific method
Scientific method refers to a body of techniques for investigating phenomena, acquiring new knowledge, or correcting and integrating previous knowledge. To be termed scientific, a method of inquiry must be based on gathering empirical and measurable evidence subject to specific principles of...
of modern science rejected Aristotle's approach, and sought explanations of natural phenomena in terms of laws of nature which were the same for all visible things, and did not need to assume any fixed natural categories, nor any divine cosmic order. But this new approach was slow to take root in the biological sciences, which became the last bastion of the concept of fixed natural types. John Ray
John Ray
John Ray was an English naturalist, sometimes referred to as the father of English natural history. Until 1670, he wrote his name as John Wray. From then on, he used 'Ray', after "having ascertained that such had been the practice of his family before him".He published important works on botany,...
used one of the previously more general terms for fixed natural types, "species", to apply to animal and plant types, but unlike Aristotle he strictly identified each type of living thing as a species, and proposed that each species can be defined by the features that perpetuate themselves each generation. These species were designed by God, but showing differences caused by local conditions. The biological classification introduced by Carolus Linnaeus
Carolus Linnaeus
Carl Linnaeus , also known after his ennoblement as , was a Swedish botanist, physician, and zoologist, who laid the foundations for the modern scheme of binomial nomenclature. He is known as the father of modern taxonomy, and is also considered one of the fathers of modern ecology...
in 1735 also viewed species as fixed according to a divine plan.
Other naturalists of this time speculated on evolutionary change of species over time according to natural laws. Maupertuis
Maupertuis
Maupertuis may refer to:*Pierre Louis Maupertuis , a French mathematician and philosopher*Maupertuis, Manche, a commune in Manche, France*Maleperduis , the castle of Reynard in the Reynard cycle*Maupertus-sur-Mer, a commune in Manche, France...
wrote in 1751 of natural modifications occurring during reproduction and accumulating over many generations to produce new species. Buffon
Georges-Louis Leclerc, Comte de Buffon
Georges-Louis Leclerc, Comte de Buffon was a French naturalist, mathematician, cosmologist, and encyclopedic author.His works influenced the next two generations of naturalists, including Jean-Baptiste Lamarck and Georges Cuvier...
suggested that species could degenerate into different organisms, and Erasmus Darwin
Erasmus Darwin
Erasmus Darwin was an English physician who turned down George III's invitation to be a physician to the King. One of the key thinkers of the Midlands Enlightenment, he was also a natural philosopher, physiologist, slave trade abolitionist,inventor and poet...
proposed that all warm-blooded animals could have descended from a single micro-organism (or "filament"). The first fully-fledged evolutionary scheme was Lamarck's "transmutation" theory of 1809 which envisaged spontaneous generation continually producing simple forms of life developed greater complexity in parallel lineages with an inherent progressive tendency, and that on a local level these lineages adapted to the environment by inheriting changes caused by use or disuse in parents. (The latter process was later called Lamarckism.) These ideas were condemned by establishment naturalists as speculation lacking empirical support. In particular Georges Cuvier
Georges Cuvier
Georges Chrétien Léopold Dagobert Cuvier or Jean Léopold Nicolas Frédéric Cuvier , known as Georges Cuvier, was a French naturalist and zoologist...
insisted that species were unrelated and fixed, their similarities reflecting divine design for functional needs. In the meantime, Ray's ideas of benevolent design had been developed by William Paley
William Paley
William Paley was a British Christian apologist, philosopher, and utilitarian. He is best known for his exposition of the teleological argument for the existence of God in his work Natural Theology, which made use of the watchmaker analogy .-Life:Paley was Born in Peterborough, England, and was...
into a natural theology
Natural theology
Natural theology is a branch of theology based on reason and ordinary experience. Thus it is distinguished from revealed theology which is based on scripture and religious experiences of various kinds; and also from transcendental theology, theology from a priori reasoning.Marcus Terentius Varro ...
which proposed complex adaptations as evidence of divine design, and was admired by Charles Darwin.
The critical break from the concept of fixed species in biology began with the theory of evolution by natural selection, which was formulated by Charles Darwin. Partly influenced by An Essay on the Principle of Population
An Essay on the Principle of Population
The book An Essay on the Principle of Population was first published anonymously in 1798 through J. Johnson . The author was soon identified as The Reverend Thomas Robert Malthus. While it was not the first book on population, it has been acknowledged as the most influential work of its era...
by Thomas Robert Malthus, Darwin noted that population growth would lead to a "struggle for existence" where favorable variations could prevail as others perished. Each generation, many offspring fail to survive to an age of reproduction because of limited resources. This could explain the diversity of animals and plants from a common ancestry through the working of natural laws working the same for all types of thing. Darwin was developing his theory of "natural selection
Natural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....
" from 1838 onwards until Alfred Russel Wallace
Alfred Russel Wallace
Alfred Russel Wallace, OM, FRS was a British naturalist, explorer, geographer, anthropologist and biologist...
sent him a similar theory in 1858. Both men presented their separate papers to the Linnean Society of London
Linnean Society of London
The Linnean Society of London is the world's premier society for the study and dissemination of taxonomy and natural history. It publishes a zoological journal, as well as botanical and biological journals...
. At the end of 1859, Darwin's publication of On the Origin of Species explained natural selection in detail and in a way that lead to an increasingly wide acceptance of Darwinian evolution
Darwinism
Darwinism is a set of movements and concepts related to ideas of transmutation of species or of evolution, including some ideas with no connection to the work of Charles Darwin....
. Thomas Henry Huxley applied Darwin's ideas to humans, using paleontology
Paleontology
Paleontology "old, ancient", ὄν, ὀντ- "being, creature", and λόγος "speech, thought") is the study of prehistoric life. It includes the study of fossils to determine organisms' evolution and interactions with each other and their environments...
and comparative anatomy
Comparative anatomy
Comparative anatomy is the study of similarities and differences in the anatomy of organisms. It is closely related to evolutionary biology and phylogeny .-Description:...
to provide strong evidence that humans and apes shared a common ancestry. Some were disturbed by this since it implied that humans did not have a special place in the universe.
Precise mechanisms of reproductive heritability and the origin of new traits remained a mystery. Towards this end, Darwin developed his provisional theory of pangenesis
Pangenesis
Pangenesis was Charles Darwin's hypothetical mechanism for heredity. He presented this 'provisional hypothesis' in his 1868 work The Variation of Animals and Plants under Domestication and felt that it brought 'together a multitude of facts which are at present left disconnected by any efficient...
. In 1865 Gregor Mendel
Gregor Mendel
Gregor Johann Mendel was an Austrian scientist and Augustinian friar who gained posthumous fame as the founder of the new science of genetics. Mendel demonstrated that the inheritance of certain traits in pea plants follows particular patterns, now referred to as the laws of Mendelian inheritance...
reported that traits were inherited in a predictable manner through the independent assortment and segregation of elements (later known as genes
Gênes
Gênes is the name of a département of the First French Empire in present Italy, named after the city of Genoa. It was formed in 1805, when Napoleon Bonaparte occupied the Republic of Genoa. Its capital was Genoa, and it was divided in the arrondissements of Genoa, Bobbio, Novi Ligure, Tortona and...
). Mendel's laws of inheritance eventually supplanted most of Darwin's pangenesis theory. August Weismann
August Weismann
Friedrich Leopold August Weismann was a German evolutionary biologist. Ernst Mayr ranked him the second most notable evolutionary theorist of the 19th century, after Charles Darwin...
made the important distinction between germ cells (sperm and eggs)
Germline
In biology and genetics, the germline of a mature or developing individual is the line of germ cells that have genetic material that may be passed to a child.For example, gametes such as the sperm or the egg, are part of the germline...
and somatic cells of the body, demonstrating that heredity passes through the germ line only. Hugo de Vries
Hugo de Vries
Hugo Marie de Vries ForMemRS was a Dutch botanist and one of the first geneticists. He is known chiefly for suggesting the concept of genes, rediscovering the laws of heredity in the 1890s while unaware of Gregor Mendel's work, for introducing the term "mutation", and for developing a mutation...
connected Darwin's pangenesis theory to Wiesman's germ/soma cell distinction and proposed that Darwin's pangenes were concentrated in the cell nucleus
Cell nucleus
In cell biology, the nucleus is a membrane-enclosed organelle found in eukaryotic cells. It contains most of the cell's genetic material, organized as multiple long linear DNA molecules in complex with a large variety of proteins, such as histones, to form chromosomes. The genes within these...
and when expressed they could move into the cytoplasm
Cytoplasm
The cytoplasm is a small gel-like substance residing between the cell membrane holding all the cell's internal sub-structures , except for the nucleus. All the contents of the cells of prokaryote organisms are contained within the cytoplasm...
to change the cells structure. De Vries was also one of the researchers who made Mendel's work well-known, believing that Mendelian traits corresponded to the transfer of heritable variations along the germline. To explain how new variants originate, De Vries developed a mutation
Mutation
In molecular biology and genetics, mutations are changes in a genomic sequence: the DNA sequence of a cell's genome or the DNA or RNA sequence of a virus. They can be defined as sudden and spontaneous changes in the cell. Mutations are caused by radiation, viruses, transposons and mutagenic...
theory that led to a temporary rift between those who accepted Darwinian evolution and biometricians who allied with de Vries. At the turn of the 20th century, pioneers in the field of population genetics, such as J.B.S. Haldane, Sewall Wright
Sewall Wright
Sewall Green Wright was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. With R. A. Fisher and J.B.S. Haldane, he was a founder of theoretical population genetics. He is the discoverer of the inbreeding coefficient and of...
, and Ronald Fisher
Ronald Fisher
Sir Ronald Aylmer Fisher FRS was an English statistician, evolutionary biologist, eugenicist and geneticist. Among other things, Fisher is well known for his contributions to statistics by creating Fisher's exact test and Fisher's equation...
, set the foundations of evolution onto a robust statistical philosophy. The false contradiction between Darwin's theory, genetic mutations, and Mendelian inheritance was thus reconciled.
In the 1920s and 1930s a modern evolutionary synthesis connected natural selection, mutation theory, and Mendelian inheritance into a unified theory that applied generally to any branch of biology. The modern synthesis was able to explain patterns observed across species in populations, through fossil transitions in palaeontology, and even complex cellular mechanisms in developmental biology
Developmental biology
Developmental biology is the study of the process by which organisms grow and develop. Modern developmental biology studies the genetic control of cell growth, differentiation and "morphogenesis", which is the process that gives rise to tissues, organs and anatomy.- Related fields of study...
. The publication of the structure of DNA by James Watson
James D. Watson
James Dewey Watson is an American molecular biologist, geneticist, and zoologist, best known as one of the co-discoverers of the structure of DNA in 1953 with Francis Crick...
and Francis Crick
Francis Crick
Francis Harry Compton Crick OM FRS was an English molecular biologist, biophysicist, and neuroscientist, and most noted for being one of two co-discoverers of the structure of the DNA molecule in 1953, together with James D. Watson...
in 1953 demonstrated a physical basis for inheritance. Molecular biology
Molecular biology
Molecular biology is the branch of biology that deals with the molecular basis of biological activity. This field overlaps with other areas of biology and chemistry, particularly genetics and biochemistry...
improved our understanding of the relationship between genotype and phenotype. Advancements were also made in phylogenetic systematics
Systematics
Biological systematics is the study of the diversification of terrestrial life, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees...
, mapping the transition of traits into a comparative and testable framework through the publication and use of evolutionary trees
Phylogenetic tree
A phylogenetic tree or evolutionary tree is a branching diagram or "tree" showing the inferred evolutionary relationships among various biological species or other entities based upon similarities and differences in their physical and/or genetic characteristics...
. In 1973, evolutionary biologist Theodosius Dobzhansky
Theodosius Dobzhansky
Theodosius Grygorovych Dobzhansky ForMemRS was a prominent geneticist and evolutionary biologist, and a central figure in the field of evolutionary biology for his work in shaping the unifying modern evolutionary synthesis...
penned that "nothing in biology makes sense except in the light of evolution", because it has brought to light the relations of what first seemed disjointed facts in natural history into a coherent explanatory body of knowledge that describes and predicts many observable facts about life on this planet.
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. This extension has been dubbed "eco-evo-devo
Evolutionary developmental biology
Evolutionary developmental biology is a field of biology that compares the developmental processes of different organisms to determine the ancestral relationship between them, and to discover how developmental processes evolved...
".
Heredity
Evolution in organisms occurs through changes in heritable traitsTrait (biology)
A trait is a distinct variant of a phenotypic character of an organism that may be inherited, environmentally determined or be a combination of the two...
– particular 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. Inherited traits are controlled by gene
Gene
A gene is a molecular unit of heredity of a living organism. It is a name given to some stretches of DNA and RNA that code for a type of protein or for an RNA chain that has a function in the organism. Living beings depend on genes, as they specify all proteins and functional RNA chains...
s and the complete set of genes within an organism's genome
Genome
In modern molecular biology and genetics, the genome is the entirety of an organism's hereditary information. It is encoded either in DNA or, for many types of virus, in RNA. The genome includes both the genes and the non-coding sequences of the DNA/RNA....
is called its genotype
Genotype
The genotype is the genetic makeup of a cell, an organism, or an individual usually with reference to a specific character under consideration...
.
The complete set of observable traits that make up the structure and behaviour of an organism is called its phenotype
Phenotype
A phenotype is an organism's observable characteristics or traits: such as its morphology, development, biochemical or physiological properties, behavior, and products of behavior...
. These traits come from the interaction of its genotype with the environment
Environment (biophysical)
The biophysical environment is the combined modeling of the physical environment and the biological life forms within the environment, and includes all variables, parameters as well as conditions and modes inside the Earth's biosphere. The biophysical environment can be divided into two categories:...
. As a result, many aspects of an organism's phenotype are not inherited. For example, suntanned
Sun tanning
Sun tanning or simply tanning is the process whereby skin color is darkened or tanned. The process is most often a result of exposure to ultraviolet radiation from the sun or from artificial sources, such as a tanning bed, but can also be a result of windburn or reflected light...
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 their genotype; a striking example are people with the inherited trait of albinism
Albinism
Albinism is a congenital disorder characterized by the complete or partial absence of pigment in the skin, hair and eyes due to absence or defect of an enzyme involved in the production of melanin...
, who do not tan at all and are very sensitive to sunburn
Sunburn
A sunburn is a burn to living tissue, such as skin, which is produced by overexposure to ultraviolet radiation, commonly from the sun's rays. Usual mild symptoms in humans and other animals include red or reddish skin that is hot to the touch, general fatigue, and mild dizziness. An excess of UV...
.
Heritable traits are known to be passed from one generation to the next via DNA
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...
, a molecule
Molecule
A molecule is an electrically neutral group of at least two atoms held together by covalent chemical bonds. Molecules are distinguished from ions by their electrical charge...
that encodes genetic information. DNA is a long polymer
Polymer
A polymer is a large molecule composed of repeating structural units. These subunits are typically connected by covalent chemical bonds...
composed of four types of bases. 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. Before a cell divides, the DNA is copied, so that each of the resulting two cells will inherit the DNA sequence. Portions of a DNA molecule that specify a single functional unit are called gene
Gene
A gene is a molecular unit of heredity of a living organism. It is a name given to some stretches of DNA and RNA that code for a type of protein or for an RNA chain that has a function in the organism. Living beings depend on genes, as they specify all proteins and functional RNA chains...
s; different genes have different sequences of bases. Within cells
Cell (biology)
The cell is the basic structural and functional unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often called the building block of life. The Alberts text discusses how the "cellular building blocks" move to shape developing embryos....
, the long strands of DNA form condensed structures called chromosome
Chromosome
A chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...
s. The specific location of a DNA sequence within a chromosome is known as a locus
Locus (genetics)
In the fields of genetics and genetic computation, a locus is the specific location of a gene or DNA sequence on a chromosome. A variant of the DNA sequence at a given locus is called an allele. The ordered list of loci known for a particular genome is called a genetic map...
. If the DNA sequence at a locus varies between individuals, the different forms of this sequence are called allele
Allele
An allele is one of two or more forms of a gene or a genetic locus . "Allel" is an abbreviation of allelomorph. Sometimes, different alleles can result in different observable phenotypic traits, such as different pigmentation...
s. DNA sequences can change through mutation
Mutation
In molecular biology and genetics, mutations are changes in a genomic sequence: the DNA sequence of a cell's genome or the DNA or RNA sequence of a virus. They can be defined as sudden and spontaneous changes in the cell. Mutations are caused by radiation, viruses, transposons and mutagenic...
s, 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. However, while this simple correspondence between an allele and a trait works in some cases, most traits are more complex and are controlled by multiple interacting genes
Quantitative trait locus
Quantitative traits refer to phenotypes that vary in degree and can be attributed to polygenic effects, i.e., product of two or more genes, and their environment. Quantitative trait loci are stretches of DNA containing or linked to the genes that underlie a quantitative trait...
.
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. DNA methylation
DNA methylation
DNA methylation is a biochemical process that is important for normal development in higher organisms. It involves the addition of a methyl group to the 5 position of the cytosine pyrimidine ring or the number 6 nitrogen of the adenine purine ring...
marking chromatin
Chromatin
Chromatin is the combination of DNA and proteins that make up the contents of the nucleus of a cell. The primary functions of chromatin are; to package DNA into a smaller volume to fit in the cell, to strengthen the DNA to allow mitosis and meiosis and prevent DNA damage, and to control gene...
, self-sustaining metabolic loops, gene silencing by RNA interference
RNA interference
RNA interference is a process within living cells that moderates the activity of their genes. Historically, it was known by other names, including co-suppression, post transcriptional gene silencing , and quelling. Only after these apparently unrelated processes were fully understood did it become...
and the three dimensional conformation
Protein structure
Proteins are an important class of biological macromolecules present in all organisms. Proteins are polymers of amino acids. Classified by their physical size, proteins are nanoparticles . Each protein polymer – also known as a polypeptide – consists of a sequence formed from 20 possible L-α-amino...
of proteins (such as prions) are areas where epigenetic inheritance systems have been discovered at the organismic level. Developmental biologists suggest that complex interactions in genetic networks
Gene regulatory network
A gene regulatory network or genetic regulatory network is a collection of DNA segments in a cell whichinteract with each other indirectly and with other substances in the cell, thereby governing the rates at which genes in the network are transcribed into mRNA.In general, each mRNA molecule goes...
and communication among cells can lead to heritable variations that may underlay some of the mechanics in developmental plasticity
Developmental plasticity
Developmental plasticity is a general term referring to changes in neural connections during development as a result of environmental interactions as well as neural changes induced by learning...
and canalization
Canalisation (genetics)
Canalisation is a measure of the ability of a population to produce the same phenotype regardless of variability of its environment or genotype. In other words, it means robustness. The term canalisation was coined by C. H. Waddington...
. Heritability may also occur at even larger scales. For example, ecological inheritance through the process of niche construction
Niche construction
Niche construction is the process in which an organism alters its own environment, often but not always in a manner that increases its chances of survival...
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. Other examples of heritability in evolution that are not under the direct control of genes include the inheritance of cultural traits
Dual inheritance theory
Dual inheritance theory , also known as gene-culture coevolution, was developed in the late 1970s and early 1980s to explain how human behavior is a product of two different and interacting evolutionary processes: genetic evolution and cultural evolution...
and symbiogenesis
Symbiogenesis
Symbiogenesis is the merging of two separate organisms to form a single new organism. The idea originated with Konstantin Mereschkowsky in his 1926 book Symbiogenesis and the Origin of Species, which proposed that chloroplasts originate from cyanobacteria captured by a protozoan...
.
Variation
An individual organism's phenotypePhenotype
A phenotype is an organism's observable characteristics or traits: such as its morphology, development, biochemical or physiological properties, behavior, and products of behavior...
results from both its genotype
Genotype
The genotype is the genetic makeup of a cell, an organism, or an individual usually with reference to a specific character under consideration...
and the influence from the environment
Environment (biophysical)
The biophysical environment is the combined modeling of the physical environment and the biological life forms within the environment, and includes all variables, parameters as well as conditions and modes inside the Earth's biosphere. The biophysical environment can be divided into two categories:...
it has lived in. A substantial part of the variation in phenotypes in a population is caused by the differences between their genotypes. The modern evolutionary synthesis
Modern evolutionary synthesis
The modern evolutionary synthesis is a union of ideas from several biological specialties which provides a widely accepted account of evolution...
defines evolution as the change over time in this genetic 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
Fixation (population genetics)
In population genetics, fixation is the change in a gene pool from a situation where there exist at least two variants of a particular gene to a situation where only one of the alleles remains...
— 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
Genetic variation
Genetic variation, variation in alleles of genes, occurs both within and among populations. Genetic variation is important because it provides the “raw material” for natural selection. Genetic variation is brought about by mutation, a change in a chemical structure of a gene. Polyploidy is an...
in a population. Before the discovery of Mendelian genetics, one common hypothesis was blending inheritance
Blending inheritance
Many biologists and other academics held to the idea of blending inheritance during the 19th century, prior to the discovery of genetics. Blending inheritance was merely a widespread hypothetical model, rather than a formalized scientific theory , in...
. But with blending inheritance, genetic variance would be rapidly lost, making evolution by natural selection implausible. The Hardy-Weinberg principle
Hardy-Weinberg principle
The Hardy–Weinberg principle states that both allele and genotype frequencies in a population remain constant—that is, they are in equilibrium—from generation to generation unless specific disturbing influences are introduced...
provides the solution to how variation is maintained in a population with Mendelian inheritance
Mendelian inheritance
Mendelian inheritance is a scientific description of how hereditary characteristics are passed from parent organisms to their offspring; it underlies much of genetics...
. The frequencies of alleles (variations in a gene) will remain constant in the absence of selection, mutation, migration and genetic drift.
Variation comes from mutation
Mutation
In molecular biology and genetics, mutations are changes in a genomic sequence: the DNA sequence of a cell's genome or the DNA or RNA sequence of a virus. They can be defined as sudden and spontaneous changes in the cell. Mutations are caused by radiation, viruses, transposons and mutagenic...
s in genetic material, reshuffling of genes through sexual reproduction
Sexual reproduction
Sexual reproduction is the creation of a new organism by combining the genetic material of two organisms. There are two main processes during sexual reproduction; they are: meiosis, involving the halving of the number of chromosomes; and fertilization, involving the fusion of two gametes and the...
and migration between populations (gene flow
Gene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...
). Despite the constant introduction of new variation through mutation and gene flow, most of the genome
Genome
In modern molecular biology and genetics, the genome is the entirety of an organism's hereditary information. It is encoded either in DNA or, for many types of virus, in RNA. The genome includes both the genes and the non-coding sequences of the DNA/RNA....
of a species is identical in all individuals of that species. However, even relatively small differences in genotype can lead to dramatic differences in phenotype: for example, chimpanzees and humans differ in only about 5% of their genomes.
Mutation
Mutations are changes in the DNA sequence of a cell's genome. When mutations occur, they can either have no effect, alter the product of a geneGene product
A gene product is the biochemical material, either RNA or protein, resulting from expression of a gene. A measurement of the amount of gene product is sometimes used to infer how active a gene is. Abnormal amounts of gene product can be correlated with disease-causing alleles, such as the...
, or prevent the gene from functioning. Based on studies in the fly Drosophila melanogaster
Drosophila melanogaster
Drosophila melanogaster is a species of Diptera, or the order of flies, in the family Drosophilidae. The species is known generally as the common fruit fly or vinegar fly. Starting from Charles W...
, it has been suggested that if a mutation changes a protein produced by a gene, this will probably be harmful, with about 70% of these mutations having damaging effects, and the remainder being either neutral or weakly beneficial.
Mutations can involve large sections of a chromosome becoming duplicated
Gene duplication
Gene duplication is any duplication of a region of DNA that contains a gene; it may occur as an error in homologous recombination, a retrotransposition event, or duplication of an entire chromosome.The second copy of the gene is often free from selective pressure — that is, mutations of it have no...
(usually by genetic recombination
Genetic recombination
Genetic recombination is a process by which a molecule of nucleic acid is broken and then joined to a different one. Recombination can occur between similar molecules of DNA, as in homologous recombination, or dissimilar molecules, as in non-homologous end joining. Recombination is a common method...
), which can introduce extra copies of a gene into a genome. Extra copies of genes are a major source of the raw material needed for new genes to evolve. This is important because most new genes evolve within gene families
Gene family
A gene family is a set of several similar genes, formed by duplication of a single original gene, and generally with similar biochemical functions...
from pre-existing genes that share common ancestors. For example, the human eye uses four genes to make structures that sense light: three for colour vision
Cone cell
Cone cells, or cones, are photoreceptor cells in the retina of the eye that are responsible for color vision; they function best in relatively bright light, as opposed to rod cells that work better in dim light. If the retina is exposed to an intense visual stimulus, a negative afterimage will be...
and one for night vision
Rod cell
Rod cells, or rods, are photoreceptor cells in the retina of the eye that can function in less intense light than can the other type of visual photoreceptor, cone cells. Named for their cylindrical shape, rods are concentrated at the outer edges of the retina and are used in peripheral vision. On...
; all four are descended from a single ancestral gene.
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
Redundancy (engineering)
In engineering, redundancy is the duplication of critical components or functions of a system with the intention of increasing reliability of the system, usually in the case of a backup or fail-safe....
of the system; one gene in the pair can acquire a new function while the other copy continues to perform its original function. Other types of mutations can even generate entirely new genes from previously noncoding DNA.
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. When new genes are assembled from shuffling pre-existing parts, domains
Protein domain
A protein domain is a part of protein sequence and structure that can evolve, function, and exist independently of the rest of the protein chain. Each domain forms a compact three-dimensional structure and often can be independently stable and folded. Many proteins consist of several structural...
act as modules with simple independent functions, which can be mixed together to produce new combinations with new and complex functions. For example, polyketide synthase
Polyketide synthase
Polyketide synthases are a family of multi-domain enzymes or enzyme complexes that produce polyketides, a large class of secondary metabolites, in bacteria, fungi, plants, and a few animal lineages...
s are large enzymes that make antibiotics; they contain up to one hundred independent domains that each catalyze one step in the overall process, like a step in an assembly line.
Sex and recombination
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 sexSex
In biology, sex is a process of combining and mixing genetic traits, often resulting in the specialization of organisms into a male or female variety . Sexual reproduction involves combining specialized cells to form offspring that inherit traits from both parents...
ual organisms contain random mixtures of their parents' chromosomes that are produced through independent assortment. In a related process called homologous recombination
Homologous recombination
Homologous recombination is a type of genetic recombination in which nucleotide sequences are exchanged between two similar or identical molecules of DNA. It is most widely used by cells to accurately repair harmful breaks that occur on both strands of DNA, known as double-strand breaks...
, sexual organisms exchange DNA between two matching chromosomes. Recombination and reassortment do not alter allele frequencies, but instead change which alleles are associated with each other, producing offspring with new combinations of alleles. Sex usually increases genetic variation and may increase the rate of evolution.
Gene flow
Gene flowGene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...
is the exchange of genes between populations and between species. It can therefore be a source of variation that is new to a population or to a species. Gene flow
Gene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...
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
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...
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
Horizontal gene transfer , also lateral gene transfer , is any process in which an organism incorporates genetic material from another organism without being the offspring of that organism...
. Horizontal gene transfer
Horizontal gene transfer
Horizontal gene transfer , also lateral gene transfer , is any process in which an organism incorporates genetic material from another organism without being the offspring of that organism...
is the transfer of genetic material from one organism to another organism that is not its offspring; this is most common among bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
. In medicine, this contributes to the spread of antibiotic resistance
Antibiotic resistance
Antibiotic resistance is a type of drug resistance where a microorganism is able to survive exposure to an antibiotic. While a spontaneous or induced genetic mutation in bacteria may confer resistance to antimicrobial drugs, genes that confer resistance can be transferred between bacteria in a...
, as when one bacteria acquires resistance genes it can rapidly transfer them to other species. Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces cerevisiae
Saccharomyces cerevisiae
Saccharomyces cerevisiae is a species of yeast. It is perhaps the most useful yeast, having been instrumental to baking and brewing since ancient times. It is believed that it was originally isolated from the skin of grapes...
and the adzuki bean beetle Callosobruchus chinensis has occurred. An example of larger-scale transfers are the eukaryotic bdelloid rotifers, which have received a range of genes from bacteria, fungi and plants. Virus
Virus
A virus is a small infectious agent that can replicate only inside the living cells of organisms. Viruses infect all types of organisms, from animals and plants to bacteria and archaea...
es can also carry DNA between organisms, allowing transfer of genes even across biological domains
Domain (biology)
In biological taxonomy, a domain is the highest taxonomic rank of organisms, higher than a kingdom. According to the three-domain system of Carl Woese, introduced in 1990, the Tree of Life consists of three domains: Archaea, Bacteria and Eukarya...
.
Large-scale gene transfer has also occurred between the ancestors of eukaryotic cells
Eukaryote
A eukaryote is an organism whose cells contain complex structures enclosed within membranes. Eukaryotes may more formally be referred to as the taxon Eukarya or Eukaryota. The defining membrane-bound structure that sets eukaryotic cells apart from prokaryotic cells is the nucleus, or nuclear...
and bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
, during the acquisition of chloroplast
Chloroplast
Chloroplasts are organelles found in plant cells and other eukaryotic organisms that conduct photosynthesis. Chloroplasts capture light energy to conserve free energy in the form of ATP and reduce NADP to NADPH through a complex set of processes called photosynthesis.Chloroplasts are green...
s and mitochondria
Mitochondrion
In cell biology, a mitochondrion is a membrane-enclosed organelle found in most eukaryotic cells. These organelles range from 0.5 to 1.0 micrometers in diameter...
. It is possible that eukaryotes themselves originated from horizontal gene transfers between bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
and archaea
Archaea
The Archaea are a group of single-celled microorganisms. A single individual or species from this domain is called an archaeon...
.
Mechanisms
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 selectionNatural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....
, genetic drift
Genetic drift
Genetic drift or allelic drift is the change in the frequency of a gene variant in a population due to random sampling.The alleles in the offspring are a sample of those in the parents, and chance has a role in determining whether a given individual survives and reproduces...
, genetic hitchhiking
Genetic hitchhiking
Genetic hitchhiking is the process by which an allele may increase in frequency by virtue of being linked to a gene that is positively selected. Proximity on a chromosome may allow genes to be dragged along with a selective sweep experienced by an advantageous gene nearby...
, mutation
Mutation
In molecular biology and genetics, mutations are changes in a genomic sequence: the DNA sequence of a cell's genome or the DNA or RNA sequence of a virus. They can be defined as sudden and spontaneous changes in the cell. Mutations are caused by radiation, viruses, transposons and mutagenic...
and gene flow
Gene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...
.
Natural selection
Evolution by means of natural selectionNatural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....
is the process by which genetic mutations that enhance reproduction become and remain, 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:
- Heritable variation exists within populations of organisms.
- Organisms produce more offspring than can survive.
- These offspring vary in their ability to survive and reproduce.
These conditions produce competition between organisms for survival and reproduction. Consequently, organisms with traits that give them an advantage over their competitors pass these advantageous traits on, while traits that do not confer an advantage are not passed on to the next generation.
The central concept of natural selection is the evolutionary fitness
Fitness (biology)
Fitness is a central idea in evolutionary theory. It can be defined either with respect to a genotype or to a phenotype in a given environment...
of an organism. Fitness is measured by an organism's ability to survive and reproduce, which determines the size of its genetic contribution to the next generation. However, fitness is not the same as the total number of offspring: instead fitness is indicated by the proportion of subsequent generations that carry an organism's genes. For example, if an organism could survive well and reproduce rapidly, but its offspring were all too small and weak to survive, this organism would make little genetic contribution to future generations and would thus have low fitness.
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". Examples of traits that can increase fitness are enhanced survival and increased fecundity
Fecundity
Fecundity, derived from the word fecund, generally refers to the ability to reproduce. In demography, fecundity is the potential reproductive capacity of an individual or population. In biology, the definition is more equivalent to fertility, or the actual reproductive rate of an organism or...
. Conversely, the lower fitness caused by having a less beneficial or deleterious allele results in this allele becoming rarer — they are "selected against". Importantly, the fitness of an allele is not a fixed characteristic; if the environment changes, previously neutral or harmful traits may become beneficial and previously beneficial traits become harmful. However, even if the direction of selection does reverse in this way, traits that were lost in the past may not re-evolve in an identical form (see Dollo's law
Dollo's law
Dollo's law of irreversibility is a hypothesis proposed in 1893 by French-born Belgian paleontologist Louis Dollo which states that evolution is not reversible...
).
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
Directional selection
In population genetics, directional selection is a mode of natural selection in which a single phenotype is favored, causing the allele frequency to continuously shift in one direction...
, which is a shift in the average value of a trait over time — for example, organisms slowly getting taller. Secondly, disruptive selection
Disruptive selection
Disruptive selection, also called diversifying selection, describes changes in population genetics in which extreme values for a trait are favored over intermediate values. In this case, the variance of the trait increases and the population is divided into two distinct groups...
is selection for extreme trait values and often results in two different values
Bimodal distribution
In statistics, a bimodal distribution is a continuous probability distribution with two different modes. These appear as distinct peaks in the probability density function, as shown in Figure 1....
becoming most common, with selection against the average value. This would be when either short or tall organisms had an advantage, but not those of medium height. Finally, in stabilizing selection
Stabilizing selection
-Description:Stabilizing or ambidirectional selection, , is a type of natural selection in which genetic diversity decreases as the population stabilizes on a particular trait value. This is probably the most common mechanism of action for natural selection...
there is selection against extreme trait values on both ends, which causes a decrease in variance
Variance
In probability theory and statistics, the variance is a measure of how far a set of numbers is spread out. It is one of several descriptors of a probability distribution, describing how far the numbers lie from the mean . In particular, the variance is one of the moments of a distribution...
around the average value and less diversity. This would, for example, cause organisms to slowly become all the same height.
A special case of natural selection is sexual selection
Sexual selection
Sexual selection, a concept introduced by Charles Darwin in his 1859 book On the Origin of Species, is a significant element of his theory of natural selection...
, which is selection for any trait that increases mating success by increasing the attractiveness of an organism to potential mates. Traits that evolved through sexual selection are particularly prominent in males of some animal species, despite traits such as cumbersome antlers, mating calls or bright colours that attract predators, decreasing the survival of individual males. This survival disadvantage is balanced by higher reproductive success in males that show these hard to fake
Handicap principle
The handicap principle is a hypothesis originally proposed in 1975 by biologist Amotz Zahavi to explain how evolution may lead to "honest" or reliable signaling between animals who have an obvious motivation to bluff or deceive each other...
, sexually selected traits.
Natural selection most generally makes nature the measure against which individuals and individual traits, are more or less likely to survive. "Nature" in this sense refers to an ecosystem
Ecosystem
An ecosystem is a biological environment consisting of all the organisms living in a particular area, as well as all the nonliving , physical components of the environment with which the organisms interact, such as air, soil, water and sunlight....
, that is, a system in which organisms interact with every other element, physical as well as biological
Biotic component
Biotic components are the living things that shape an ecosystem. A biotic factor is any living component that affects another organism, including animals that consume the organism in question, and the living food that the organism consumes. Each biotic factor needs energy to do work and food for...
, in their local environment
Environment (biophysical)
The biophysical environment is the combined modeling of the physical environment and the biological life forms within the environment, and includes all variables, parameters as well as conditions and modes inside the Earth's biosphere. The biophysical environment can be divided into two categories:...
. Eugene Odum, a founder of ecology, defined an ecosystem as: "Any unit that includes all of the organisms...in a given area interacting with the physical environment so that a flow of energy leads to clearly defined trophic structure, biotic diversity and material cycles (ie: exchange of materials between living and nonliving parts) within the system." Each population within an ecosystem occupies a distinct niche
Ecological niche
In ecology, a niche is a term describing the relational position of a species or population in its ecosystem to each other; e.g. a dolphin could potentially be in another ecological niche from one that travels in a different pod if the members of these pods utilize significantly different food...
, or position, with distinct relationships to other parts of the system. These relationships involve the life history of the organism, its position in the food chain
Food chain
A food web depicts feeding connections in an ecological community. Ecologists can broadly lump all life forms into one of two categories called trophic levels: 1) the autotrophs, and 2) the heterotrophs...
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
Unit of selection
A unit of selection is a biological entity within the hierarchy of biological organisation that is subject to natural selection...
, such as genes, cells, individual organisms, groups of organisms and species. Selection can act at multiple levels simultaneously. An example of selection occurring below the level of the individual organism are genes called transposon
Transposon
Transposable elements are sequences of DNA that can move or transpose themselves to new positions within the genome of a single cell. The mechanism of transposition can be either "copy and paste" or "cut and paste". Transposition can create phenotypically significant mutations and alter the cell's...
s, which can replicate and spread throughout a genome
Genome
In modern molecular biology and genetics, the genome is the entirety of an organism's hereditary information. It is encoded either in DNA or, for many types of virus, in RNA. The genome includes both the genes and the non-coding sequences of the DNA/RNA....
. Selection at a level above the individual, such as group selection
Group selection
In evolutionary biology, group selection refers to the idea that alleles can become fixed or spread in a population because of the benefits they bestow on groups, regardless of the alleles' effect on the fitness of individuals within that group....
, may allow the evolution of co-operation, as discussed below.
Biased mutation
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. If two genotypes, for example one with the nucleotide G and another with the nucleotide A in the same position, have the same fitness, but mutation from G to A happens more often than mutation from A to G, then genotypes with A will tend to evolve. Different insertion vs. deletion mutation biases in different taxa can lead to the evolution of different genome sizes. Developmental or mutational biases have also been observed in morphologicalMorphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
evolution. For example, according to the phenotype-first theory of evolution
Baldwin effect
The Baldwin effect, also known as Baldwinian evolution or ontogenic evolution, is a theory of a possible evolutionary processes that was originally put forward in 1896 in a paper, "A New Factor in Evolution," by American psychologist James Mark Baldwin. The paper proposed a mechanism for specific...
, mutations can eventually cause the genetic assimilation
Genetic assimilation
Note: Genetic assimilation is sometimes used to describe "eventual extinction of a natural species as massive pollen flow occurs from another related species and the older crop becomes more like the new crop." This usage is unrelated to the usage below....
of traits that were previously induced by the environment
Phenotypic plasticity
Phenotypic plasticity is the ability of an organism to change its phenotype in response to changes in the environment. Such plasticity in some cases expresses as several highly morphologically distinct results; in other cases, a continuous norm of reaction describes the functional interrelationship...
.
Mutation bias effects are superimposed on other processes. If selection would favor 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. Mutations leading to the loss of function of a gene are much more common than mutations that produce a new, fully functional gene. Most loss of function mutations are selected against. But when selection is weak, mutation bias towards loss of function can affect evolution. For example, pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...
s are no longer useful when animals live in the darkness of caves, and tend to be lost. This kind of loss of function can occur because of mutation bias, and/or because the function had a cost, and once the benefit of the function disappeared, natural selection leads to the loss. Loss of sporulation
Endospore
An endospore is a dormant, tough, and temporarily non-reproductive structure produced by certain bacteria from the Firmicute phylum. The name "endospore" is suggestive of a spore or seed-like form , but it is not a true spore . It is a stripped-down, dormant form to which the bacterium can reduce...
ability in a bacterium
Bacillus subtilis
Bacillus subtilis, known also as the hay bacillus or grass bacillus, is a Gram-positive, catalase-positive bacterium commonly found in soil. A member of the genus Bacillus, B. subtilis is rod-shaped, and has the ability to form a tough, protective endospore, allowing the organism to tolerate...
during laboratory evolution appears to have been caused by mutation bias, rather than natural selection against the cost of maintaining sporulation ability. When there is no selection for loss of function, the speed at which loss evolves depends more on the mutation rate than it does on the effective population size
Effective population size
In population genetics, the concept of effective population size Ne was introduced by the American geneticist Sewall Wright, who wrote two landmark papers on it...
, indicating that it is driven more by mutation bias than by genetic drift.
Genetic drift
Genetic drift is the change in allele frequencyAllele frequency
Allele frequency or Gene frequency is the proportion of all copies of a gene that is made up of a particular gene variant . In other words, it is the number of copies of a particular allele divided by the number of copies of all alleles at the genetic place in a population. It can be expressed for...
from one generation to the next that occurs because alleles are subject to sampling error
Sampling error
-Random sampling:In statistics, sampling error or estimation error is the error caused by observing a sample instead of the whole population. The sampling error can be found by subtracting the value of a parameter from the value of a statistic...
. As a result, when selective forces are absent or relatively weak, allele frequencies tend to "drift" upward or downward randomly (in a random walk
Random walk
A random walk, sometimes denoted RW, is a mathematical formalisation of a trajectory that consists of taking successive random steps. For example, the path traced by a molecule as it travels in a liquid or a gas, the search path of a foraging animal, the price of a fluctuating stock and the...
). This drift halts when an allele eventually becomes fixed
Fixation (population genetics)
In population genetics, fixation is the change in a gene pool from a situation where there exist at least two variants of a particular gene to a situation where only one of the alleles remains...
, either by disappearing from the population, or replacing the other alleles entirely. 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.
It is usually difficult to measure the relative importance of selection and neutral processes, including drift. The comparative importance of adaptive and non-adaptive forces in driving evolutionary change is an area of current research
Current research in evolutionary biology
Current research in evolutionary biology covers diverse topics, as should be expected given the centrality of evolution to understanding biology...
.
The neutral theory of molecular evolution
Neutral theory of molecular evolution
The neutral theory of molecular evolution states that the vast majority of evolutionary changes at the molecular level are caused by random drift of selectively neutral mutants . The theory was introduced by Motoo Kimura in the late 1960s and early 1970s...
proposed that most evolutionary changes are the result of the fixation of neutral mutation
Neutral mutation
In genetics, a neutral mutation is a mutation that has no effect on fitness. In other words, it is neutral with respect to natural selection.For example, some mutations in a DNA triplet or codon do not change which amino acid is introduced: this is known as a synonymous substitution. Unless the...
s by genetic drift. Hence, in this model, most genetic changes in a population are the result of constant mutation pressure and genetic drift. This form of the neutral theory is now largely abandoned, since it does not seem to fit the genetic variation seen in nature. However, a more recent and better-supported version of this model is the nearly neutral theory
Nearly neutral theory of molecular evolution
The nearly neutral theory of molecular evolution is a modification of the neutral theory of molecular evolution that accounts for slightly advantageous or deleterious mutations at the molecular level...
, where a mutation that would be neutral in a small population is not necessarily neutral in a large population. Other alternative theories propose that genetic drift is dwarfed by other stochastic forces in evolution, such as genetic hitchhiking
Genetic hitchhiking
Genetic hitchhiking is the process by which an allele may increase in frequency by virtue of being linked to a gene that is positively selected. Proximity on a chromosome may allow genes to be dragged along with a selective sweep experienced by an advantageous gene nearby...
, also known as genetic draft.
The time for a neutral allele to become fixed by genetic drift depends on population size, with fixation occurring more rapidly in smaller populations. The number of individuals in a population is not critical, but instead a measure known as the effective population size
Effective population size
In population genetics, the concept of effective population size Ne was introduced by the American geneticist Sewall Wright, who wrote two landmark papers on it...
. The effective population is usually smaller than the total population since it takes into account factors such as the level of inbreeding and the stage of the lifecycle in which the population is the smallest. The effective population size may not be the same for every gene in the same population.
Genetic hitchhiking
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 linkageGenetic linkage
Genetic linkage is the tendency of certain loci or alleles to be inherited together. Genetic loci that are physically close to one another on the same chromosome tend to stay together during meiosis, and are thus genetically linked.-Background:...
. This tendency is measured by finding how often two alleles occur together on a single chromosome compared to expectations, which is called their linkage disequilibrium
Linkage disequilibrium
In population genetics, linkage disequilibrium is the non-random association of alleles at two or more loci, not necessarily on the same chromosome. It is also referred to as to as gametic phase disequilibrium , or simply gametic disequilibrium...
. A set of alleles that is usually inherited in a group is called a haplotype
Haplotype
A haplotype in genetics is a combination of alleles at adjacent locations on the chromosome that are transmitted together...
. This can be important when one allele in a particular haplotype is strongly beneficial: natural selection can drive a selective sweep
Selective sweep
A selective sweep is the reduction or elimination of variation among the nucleotides in neighboring DNA of a mutation as the result of recent and strong positive natural selection....
that will also cause the other alleles in the haplotype to become more common in the population; this effect is called genetic hitchhiking
Genetic hitchhiking
Genetic hitchhiking is the process by which an allele may increase in frequency by virtue of being linked to a gene that is positively selected. Proximity on a chromosome may allow genes to be dragged along with a selective sweep experienced by an advantageous gene nearby...
or genetic draft. Genetic draft caused by the fact that some neutral genes are genetically linked to others that are under selection can be partially captured by an appropriate effective population size.
Gene flow
Gene flowGene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...
is the exchange of genes between populations and between species. The presence or absence of gene flow fundamentally changes the course of evolution. 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
Bateson-Dobzhansky-Muller Model
The Bateson-Dobzhansky-Muller Model is a model of the evolution of genetic incompatibility. The theory was first described by William Bateson in 1909 , then independently described by Theodosius Dobzhansky in 1934 , and later elaborated by Herman Muller....
, 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 organism within these populations to evolve 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
Sewall Wright
Sewall Green Wright was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. With R. A. Fisher and J.B.S. Haldane, he was a founder of theoretical population genetics. He is the discoverer of the inbreeding coefficient and of...
's developed his shifting balance theory
Shifting balance theory
The shifting balance theory of evolution is a view of evolution advocated by Sewall Wright that each of the four evolutionary forces is important to adaptive evolution...
that gene flow between partially isolated populations was an important aspect of adaptive evolution. However, recently there has been substantial criticism of the importance of the shifting balance theory
Shifting balance theory
The shifting balance theory of evolution is a view of evolution advocated by Sewall Wright that each of the four evolutionary forces is important to adaptive evolution...
.
Outcomes
Evolution influences every aspect of the form and behaviour of organisms. Most prominent are the specific behavioural and physical adaptationAdaptation
An adaptation in biology is a trait with a current functional role in the life history of an organism that is maintained and evolved by means of natural selection. An adaptation refers to both the current state of being adapted and to the dynamic evolutionary process that leads to the adaptation....
s 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 co-operating
Co-operation (evolution)
Co-operation or co-operative behaviours are terms used to describe behaviours by organisms which are beneficial to other organisms, and are selected for on that basis. Under this definition, altruism is a form of co-operation in which there is no direct benefit to the actor...
with each other, usually by aiding their relatives or engaging in mutually beneficial symbiosis
Symbiosis
Symbiosis is close and often long-term interaction between different biological species. In 1877 Bennett used the word symbiosis to describe the mutualistic relationship in lichens...
. 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 sometimes divided into macroevolution
Macroevolution
Macroevolution is evolution on a scale of separated gene pools. Macroevolutionary studies focus on change that occurs at or above the level of species, in contrast with microevolution, which refers to smaller evolutionary changes within a species or population.The process of speciation may fall...
, which is evolution that occurs at or above the level of species, such as extinction
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...
and speciation
Speciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...
and microevolution
Microevolution
Microevolution is the changes in allele frequencies that occur over time within a population. This change is due to four different processes: mutation, selection , gene flow, and genetic drift....
, which is smaller evolutionary changes, such as adaptations, within a species or population. In general, macroevolution is regarded as the outcome of long periods of microevolution. Thus, the distinction between micro- and macroevolution is not a fundamental one – the difference is simply the time involved. However, in macroevolution, the traits of the entire species may be important. 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 or long-term plans; realistically however, evolution has no long-term goal and does not necessarily produce greater complexity. Although complex species
Evolution of complexity
The evolution of biological complexity is an important outcome of the process of evolution. Evolution has produced some remarkably complex organisms - although the actual level of complexity is very hard to define or measure accurately in biology, with properties such as gene content, the number of...
have evolved, they occur as a side effect of the overall number of organisms increasing and simple forms of life still remain more common in the biosphere. For example, the overwhelming majority of species are microscopic prokaryote
Prokaryote
The prokaryotes are a group of organisms that lack a cell nucleus , or any other membrane-bound organelles. The organisms that have a cell nucleus are called eukaryotes. Most prokaryotes are unicellular, but a few such as myxobacteria have multicellular stages in their life cycles...
s, which form about half the world's biomass
Biomass
Biomass, as a renewable energy source, is biological material from living, or recently living organisms. As an energy source, biomass can either be used directly, or converted into other energy products such as biofuel....
despite their small size, and constitute the vast majority of Earth's biodiversity. Simple organisms have therefore been the dominant form of life on Earth throughout its history and continue to be the main form of life up to the present day, with complex life only appearing more diverse because it is more noticeable
Biased sample
In statistics, sampling bias is when a sample is collected in such a way that some members of the intended population are less likely to be included than others. It results in a biased sample, a non-random sample of a population in which all individuals, or instances, were not equally likely to...
. Indeed, the evolution of microorganism
Microorganism
A microorganism or microbe is a microscopic organism that comprises either a single cell , cell clusters, or no cell at all...
s is particularly important to modern evolutionary research
Current research in evolutionary biology
Current research in evolutionary biology covers diverse topics, as should be expected given the centrality of evolution to understanding biology...
, since their rapid reproduction allows the study of experimental evolution
Experimental evolution
In evolutionary and experimental biology, the field of experimental evolution is concerned with testing hypotheses and theories of evolution by use of controlled experiments. Evolution may be observed in the laboratory as populations adapt to new environmental conditions and/or change by such...
and the observation of evolution and adaptation in real time.
Adaptation
Adaptation is the process that makes organisms better suited to their habitatHabitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...
. Also, the term adaptation may refer to a trait
Trait (biology)
A trait is a distinct variant of a phenotypic character of an organism that may be inherited, environmentally determined or be a combination of the two...
that is important for an organism's survival. 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
Natural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....
. The following definitions are due to Theodosius Dobzhansky
Theodosius Dobzhansky
Theodosius Grygorovych Dobzhansky ForMemRS was a prominent geneticist and evolutionary biologist, and a central figure in the field of evolutionary biology for his work in shaping the unifying modern evolutionary synthesis...
.
- Adaptation is the evolutionary process whereby an organism becomes better able to live in its habitatHabitat* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...
or habitats. - Adaptedness is the state of being adapted: the degree to which an organism is able to live and reproduce in a given set of habitats.
- An adaptive trait is an aspect of the developmental pattern of the organism which enables or enhances the probability of that organism surviving and reproducing.
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
Antibiotic
An antibacterial is a compound or substance that kills or slows down the growth of bacteria.The term is often used synonymously with the term antibiotic; today, however, with increased knowledge of the causative agents of various infectious diseases, antibiotic has come to denote a broader range of...
selection, with genetic changes causing antibiotic resistance
Antibiotic resistance
Antibiotic resistance is a type of drug resistance where a microorganism is able to survive exposure to an antibiotic. While a spontaneous or induced genetic mutation in bacteria may confer resistance to antimicrobial drugs, genes that confer resistance can be transferred between bacteria in a...
by both modifying the target of the drug, or increasing the activity of transporters that pump the drug out of the cell. Other striking examples are the bacteria Escherichia coli
Escherichia coli
Escherichia coli is a Gram-negative, rod-shaped bacterium that is commonly found in the lower intestine of warm-blooded organisms . Most E. coli strains are harmless, but some serotypes can cause serious food poisoning in humans, and are occasionally responsible for product recalls...
evolving the ability to use citric acid
Citric acid
Citric acid is a weak organic acid. It is a natural preservative/conservative and is also used to add an acidic, or sour, taste to foods and soft drinks...
as a nutrient in a long-term laboratory experiment
E. coli long-term evolution experiment
The E. coli long-term evolution experiment is an ongoing study in experimental evolution led by Richard Lenski that has been tracking genetic changes in 12 initially identical populations of asexual Escherichia coli bacteria since 24 February 1988...
, Flavobacterium
Flavobacterium
Flavobacterium is a genus of Gram-negative, non-motile and motile, rod-shaped bacteria that consists of ten recognized species, as well as three newly proposed species . Flavobacteria are found in soil and fresh water in a variety of environments...
evolving a novel enzyme that allows these bacteria to grow on the by-products of nylon
Nylon
Nylon is a generic designation for a family of synthetic polymers known generically as polyamides, first produced on February 28, 1935, by Wallace Carothers at DuPont's research facility at the DuPont Experimental Station...
manufacturing, and the soil bacterium Sphingobium
Sphingobium
Sphingobium species are different from other sphingomonads in that they are commonly isolated from soil, however Sphingobium yanoikuyae was isolated from a clinical specimen...
evolving an entirely new metabolic pathway
Metabolic pathway
In biochemistry, metabolic pathways are series of chemical reactions occurring within a cell. In each pathway, a principal chemical is modified by a series of chemical reactions. Enzymes catalyze these reactions, and often require dietary minerals, vitamins, and other cofactors in order to function...
that degrades the synthetic pesticide
Pesticide
Pesticides are substances or mixture of substances intended for preventing, destroying, repelling or mitigating any pest.A pesticide may be a chemical unicycle, biological agent , antimicrobial, disinfectant or device used against any pest...
pentachlorophenol
Pentachlorophenol
Pentachlorophenol is an organochlorine compound used as a pesticide and a disinfectant. First produced in the 1930s, it is marketed under many trade names...
. An interesting but still controversial idea is that some adaptations might increase the ability of organisms to generate genetic diversity and adapt by natural selection (increasing organisms' evolvability
Evolvability
Evolvability is defined as the capacity of a system for adaptive evolution. Evolvability is the ability of a population of organisms to not merely generate genetic diversity, but to generate adaptive genetic diversity, and thereby evolve through natural selection.In order for a biological organism...
).
Homology (biology)
Homology forms the basis of organization for comparative biology. In 1843, Richard Owen defined homology as "the same organ in different animals under every variety of form and function". Organs as different as a bat's wing, a seal's flipper, a cat's paw and a human hand have a common underlying...
being adapted to function in different ways. The bones within bat
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...
wings, for example, are very similar to those in mice
Mouse
A mouse is a small mammal belonging to the order of rodents. The best known mouse species is the common house mouse . It is also a popular pet. In some places, certain kinds of field mice are also common. This rodent is eaten by large birds such as hawks and eagles...
feet and primate
Primate
A primate is a mammal of the order Primates , which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment...
hands, due to the descent of all these structures from a common mammalian ancestor. However, since all living organisms are related to some extent, even organs that appear to have little or no structural similarity, such as arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...
, squid and vertebrate eyes, or the limbs and wings of arthropods and vertebrates, can depend on a common set of homologous genes that control their assembly and function; this is called deep homology
Deep homology
In evolutionary developmental biology, the concept of deep homology is used to describe cases where growth and differentiation processes are governed by genetic mechanisms that are homologous and deeply conserved across a wide range of species....
.
During evolution, some structures may lose their original function and become vestigial structure
Vestigial structure
Vestigiality describes homologous characters of organisms that have seemingly lost all or most of their original function in a species through evolution. These may take various forms such as anatomical structures, behaviors and biochemical pathways. Some of these disappear early in embryonic...
s. Such structures may have little or no function in a current species, yet have a clear function in ancestral species, or other closely related species. Examples include pseudogene
Pseudogene
Pseudogenes are dysfunctional relatives of known genes that have lost their protein-coding ability or are otherwise no longer expressed in the cell...
s, the non-functional remains of eyes in blind cave-dwelling fish, wings in flightless birds, and the presence of hip bones in whales and snakes. Examples of vestigial structures in humans
Human vestigiality
In the context of human evolution, human vestigiality involves those characters occurring in the human species that are considered vestigial—in other words having lost all or most of their original function through evolution...
include wisdom teeth, the coccyx
Coccyx
The coccyx , commonly referred to as the tailbone, is the final segment of the vertebral column. Comprising three to five separate or fused vertebrae below the sacrum, it is attached to the sacrum by a fibrocartilaginous joint, the sacrococcygeal symphysis, which permits limited movement between...
, the vermiform appendix
Vermiform appendix
The appendix is a blind-ended tube connected to the cecum , from which it develops embryologically. The cecum is a pouchlike structure of the colon...
, and other behavioural vestiges such as goose bumps
Goose bumps
Goose bumps, also called goose flesh, goose pimples, chill bumps, chicken skin, funky spots, Dasler Bumps, chicken bumps or the medical term cutis anserina, are the bumps on a person's skin at the base of body hairs which may involuntarily develop when a person is cold or experiences strong...
and primitive reflexes
Primitive reflexes
Primitive reflexes are reflex actions originating in the central nervous system that are exhibited by normal infants but not neurologically intact adults, in response to particular stimuli. These reflexes disappear or are inhibited by the frontal lobes as a child moves through normal child...
.
However, many traits that appear to be simple adaptations are in fact exaptation
Exaptation
Exaptation, cooption, and preadaptation are related terms referring to shifts in the function of a trait during evolution. For example, a trait can evolve because it served one particular function, but subsequently it may come to serve another. Exaptations are common in both anatomy and behaviour...
s: structures originally adapted for one function, but which coincidentally became somewhat useful for some other function in the process. One example is the African lizard Holaspis guentheri, which developed an extremely flat head for hiding in crevices, as can be seen by looking at its near relatives. However, in this species, the head has become so flattened that it assists in gliding from tree to tree—an exaptation
Exaptation
Exaptation, cooption, and preadaptation are related terms referring to shifts in the function of a trait during evolution. For example, a trait can evolve because it served one particular function, but subsequently it may come to serve another. Exaptations are common in both anatomy and behaviour...
. Within cells, molecular machine
Molecular machine
A molecular machine, or nanomachine, is any discrete number of molecular components that produce quasi-mechanical movements in response to specific stimuli . The expression is often more generally applied to molecules that simply mimic functions that occur at the macroscopic level...
s such as the bacterial flagella and protein sorting machinery
Translocase of the inner membrane
The translocase of the inner membrane is a complex of proteins found in the inner mitochondrial membrane of the mitochondria. Components of the TIM complex facilitate the translocation of proteins across the inner membrane and into the matrix...
evolved by the recruitment of several pre-existing proteins that previously had different functions. Another example is the recruitment of enzymes from glycolysis
Glycolysis
Glycolysis is the metabolic pathway that converts glucose C6H12O6, into pyruvate, CH3COCOO− + H+...
and xenobiotic metabolism
Xenobiotic metabolism
Xenobiotic metabolism is the set of metabolic pathways that modify the chemical structure of xenobiotics, which are compounds foreign to an organism's normal biochemistry, such as drugs and poisons...
to serve as structural proteins called crystallin
Crystallin
In anatomy, a crystallin is a water-soluble structural protein found in the lens and the cornea of the eye accounting for the transparency of the structure. It has also been identified in other places such as the heart, and in aggressive breast cancer tumors....
s within the lenses of organisms' eye
Eye
Eyes are organs that detect light and convert it into electro-chemical impulses in neurons. The simplest photoreceptors in conscious vision connect light to movement...
s.
A critical principle of ecology
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...
is that of competitive exclusion
Competitive exclusion principle
In ecology, the competitive exclusion principle, sometimes referred to as Gause's law of competitive exclusion or just Gause's law, is a proposition which states that two species competing for the same resources cannot coexist if other ecological factors are constant...
: no two species can occupy the same niche in the same environment for a long time. Consequently, natural selection will tend to force species to adapt to different ecological niche
Ecological niche
In ecology, a niche is a term describing the relational position of a species or population in its ecosystem to each other; e.g. a dolphin could potentially be in another ecological niche from one that travels in a different pod if the members of these pods utilize significantly different food...
s. This may mean that, for example, two species of cichlid
Cichlid
Cichlids are fishes from the family Cichlidae in the order Perciformes. Cichlids are members of a group known as the Labroidei along with the wrasses , damselfish , and surfperches . This family is both large and diverse. At least 1,300 species have been scientifically described, making it one of...
fish adapt to live in different habitat
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...
s, which will minimise the competition between them for food.
An area of current investigation in evolutionary developmental biology
Evolutionary developmental biology
Evolutionary developmental biology is a field of biology that compares the developmental processes of different organisms to determine the ancestral relationship between them, and to discover how developmental processes evolved...
is the developmental
Developmental biology
Developmental biology is the study of the process by which organisms grow and develop. Modern developmental biology studies the genetic control of cell growth, differentiation and "morphogenesis", which is the process that gives rise to tissues, organs and anatomy.- Related fields of study...
basis of adaptations and exaptations. This research addresses the origin and evolution of embryonic development
Embryogenesis
Embryogenesis is the process by which the embryo is formed and develops, until it develops into a fetus.Embryogenesis starts with the fertilization of the ovum by sperm. The fertilized ovum is referred to as a zygote...
and how modifications of development and developmental processes produce novel features. These studies have shown that evolution can alter development to produce new structures, such as embryonic bone structures that develop into the jaw in other animals instead forming part of the middle ear in mammals. It is also possible for structures that have been lost in evolution to reappear due to changes in developmental genes, such as a mutation in chicken
Chicken
The chicken is a domesticated fowl, a subspecies of the Red Junglefowl. As one of the most common and widespread domestic animals, and with a population of more than 24 billion in 2003, there are more chickens in the world than any other species of bird...
s causing embryos to grow teeth similar to those of crocodile
Crocodile
A crocodile is any species belonging to the family Crocodylidae . The term can also be used more loosely to include all extant members of the order Crocodilia: i.e...
s. It is now becoming clear that most alterations in the form of organisms are due to changes in a small set of conserved genes.
Co-evolution
Interactions between organisms can produce both conflict and co-operation. When the interaction is between pairs of species, such as a pathogenPathogen
A pathogen gignomai "I give birth to") or infectious agent — colloquially, a germ — is a microbe or microorganism such as a virus, bacterium, prion, or fungus that causes disease in its animal or plant host...
and a host
Host (biology)
In biology, a host is an organism that harbors a parasite, or a mutual or commensal symbiont, typically providing nourishment and shelter. In botany, a host plant is one that supplies food resources and substrate for certain insects or other fauna...
, or a predator
Predation
In ecology, predation describes a biological interaction where a predator feeds on its prey . Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of its prey and the eventual absorption of the prey's tissue through consumption...
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 co-evolution
Co-evolution
In biology, coevolution is "the change of a biological object triggered by the change of a related object." Coevolution can occur at many biological levels: it can be as microscopic as correlated mutations between amino acids in a protein, or as macroscopic as covarying traits between different...
. An example is the production of tetrodotoxin
Tetrodotoxin
Tetrodotoxin, also known as "tetrodox" and frequently abbreviated as TTX, sometimes colloquially referred to as "zombie powder" by those who practice Vodou, is a potent neurotoxin with no known antidote. There have been successful tests of a possible antidote in mice, but further tests must be...
in the rough-skinned newt
Rough-skinned Newt
The rough-skinned newt is a North American newt known for its strong poison.- Toxicity :Many newts produce toxins to avoid predation, but the toxins of the genus Taricha are particularly potent...
and the evolution of tetrodotoxin resistance in its predator, the common garter snake
Common Garter Snake
The Common Garter Snake is a snake indigenous to North America. Most garter snakes have a pattern of yellow stripes on a brown or green background and their average length is about , maximum about .-Subspecies:...
. In this predator-prey pair, an evolutionary arms race
Evolutionary arms race
In evolutionary biology, an evolutionary arms race is an evolutionary struggle between competing sets of co-evolving genes that develop adaptations and counter-adaptations against each other, resembling an arms race, which are also examples of positive feedback...
has produced high levels of toxin in the newt and correspondingly high levels of toxin resistance in the snake.
Co-operation
Not all co-evolved interactions between species involve conflict. Many cases of mutually beneficial interactions have evolved. For instance, an extreme cooperation exists between plants and the mycorrhizal fungiMycorrhiza
A mycorrhiza is a symbiotic association between a fungus and the roots of a vascular plant....
that grow on their roots and aid the plant in absorbing nutrients from the soil. This is a reciprocal
Reciprocity (evolution)
Reciprocity in evolutionary biology refers to mechanisms whereby the evolution of cooperative or altruistic behaviour may be favoured by the probability of future mutual interactions...
relationship as the plants provide the fungi with sugars from photosynthesis. Here, the fungi actually grow inside plant cells, allowing them to exchange nutrients with their hosts, while sending signals
Signal transduction
Signal transduction occurs when an extracellular signaling molecule activates a cell surface receptor. In turn, this receptor alters intracellular molecules creating a response...
that suppress the plant immune system
Immune system
An immune system is a system of biological structures and processes within an organism that protects against disease by identifying and killing pathogens and tumor cells. It detects a wide variety of agents, from viruses to parasitic worms, and needs to distinguish them from the organism's own...
.
Coalitions between organisms of the same species have also evolved. An extreme case is the eusociality
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....
found in social insects
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....
, such as bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...
s, termite
Termite
Termites are a group of eusocial insects that, until recently, were classified at the taxonomic rank of order Isoptera , but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea...
s and ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...
s, where sterile insects feed and guard the small number of organisms in a colony
Colony (biology)
In biology, a colony reference to several individual organisms of the same species living closely together, usually for mutual benefit, such as stronger defense or the ability to attack bigger prey. Some insects live only in colonies...
that are able to reproduce. On an even smaller scale, the somatic cell
Somatic cell
A somatic cell is any biological cell forming the body of an organism; that is, in a multicellular organism, any cell other than a gamete, germ cell, gametocyte or undifferentiated stem cell...
s 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 cell
Germ cell
A germ cell is any biological cell that gives rise to the gametes of an organism that reproduces sexually. In many animals, the germ cells originate near the gut of an embryo and migrate to the developing gonads. There, they undergo cell division of two types, mitosis and meiosis, followed by...
s 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
Carcinogenesis
Carcinogenesis or oncogenesis is literally the creation of cancer. It is a process by which normal cells are transformed into cancer cells...
.
Such cooperation within species may have evolved through the process of kin selection
Kin selection
Kin selection refers to apparent strategies in evolution that favor the reproductive success of an organism's relatives, even at a cost to the organism's own survival and reproduction. Charles Darwin was the first to discuss the concept of group/kin selection...
, which is where one organism acts to help raise a relative's offspring. This activity is selected for because if the helping individual contains alleles which promote the helping activity, it is likely that its kin will also contain these alleles and thus those alleles will be passed on. Other processes that may promote cooperation include group selection
Group selection
In evolutionary biology, group selection refers to the idea that alleles can become fixed or spread in a population because of the benefits they bestow on groups, regardless of the alleles' effect on the fitness of individuals within that group....
, where cooperation provides benefits to a group of organisms.
Speciation
SpeciationSpeciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...
is the process where a species diverges into two or more descendant species.
There are multiple ways to define the concept of "species". The choice of definition is dependent on the particularities of the species concerned. For example, some species concepts apply more readily toward sexually reproducing organisms while others lend themselves better toward asexual organisms. Despite the diversity of various species concepts, these various concepts can be placed into one of three broad philosophical approaches: interbreeding, ecological and phylogenetic. The biological species concept (BSC) is a classic example of the interbreeding approach. Defined by Ernst Mayr in 1942, the BSC states that "species are groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups". Despite its wide and long-term use, the BSC like others is not without controversy, for example because these concepts cannot be applied to prokaryotes, and this is called the species problem
Species problem
The species problem or species concept is a mixture of difficult, related questions that often come up when biologists identify species and when they define the word "species"....
. Some researchers have attempted a unifying monistic definition of species, while others adopt a pluralistic approach and suggest that there may be a different ways to logically interpret the definition of a species. "
Barriers to reproduction
Reproductive isolation
The mechanisms of reproductive isolation or hybridization barriers are a collection of mechanisms, behaviors and physiological processes that prevent the members of two different species that cross or mate from producing offspring, or which ensure that any offspring that may be produced is not...
between two diverging sexual populations are required for the populations to become new species
Speciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...
. 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
Most recent common ancestor
In genetics, the most recent common ancestor of any set of organisms is the most recent individual from which all organisms in the group are directly descended...
, it may still be possible for them to produce offspring, as with horse
Horse
The horse is one of two extant subspecies of Equus ferus, or the wild horse. It is a single-hooved mammal belonging to the taxonomic family Equidae. The horse has evolved over the past 45 to 55 million years from a small multi-toed creature into the large, single-toed animal of today...
s and donkey
Donkey
The donkey or ass, Equus africanus asinus, is a domesticated member of the Equidae or horse family. The wild ancestor of the donkey is the African Wild Ass, E...
s mating to produce mule
Mule
A mule is the offspring of a male donkey and a female horse. Horses and donkeys are different species, with different numbers of chromosomes. Of the two F1 hybrids between these two species, a mule is easier to obtain than a hinny...
s. Such hybrids are generally infertile
Infertility
Infertility primarily refers to the biological inability of a person to contribute to conception. Infertility may also refer to the state of a woman who is unable to carry a pregnancy to full term...
. 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. The importance of hybridisation in producing new species
Hybrid speciation
Hybrid speciation is the process wherein hybridization between two different closely related species leads to a distinct phenotype. This phenotype in very rare cases can also be fitter than the parental lineage and as such natural selection may then favor these individuals. Eventually, if...
of animals is unclear, although cases have been seen in many types of animals, with the gray tree frog being a particularly well-studied example.
Speciation has been observed multiple times under both controlled laboratory conditions and in nature. In sexually reproducing organisms, speciation results from reproductive isolation followed by genealogical divergence. There are four mechanisms for speciation. The most common in animals is allopatric speciation
Allopatric speciation
Allopatric speciation or geographic speciation is speciation that occurs when biological populations of the same species become isolated due to geographical changes such as mountain building or social changes such as emigration...
, which occurs in populations initially isolated geographically, such as by habitat fragmentation
Habitat fragmentation
Habitat fragmentation as the name implies, describes the emergence of discontinuities in an organism's preferred environment , causing population fragmentation...
or migration. Selection under these conditions can produce very rapid changes in the appearance and behaviour of organisms. As selection and drift act independently on populations isolated from the rest of their species, separation may eventually produce organisms that cannot interbreed.
The second mechanism of speciation is peripatric speciation
Peripatric speciation
Peripatric and peripatry are terms from biogeography, referring to organisms whose ranges are closely adjacent but do not overlap, being separated where these organisms do not occur – for example a wide river or a mountain range. Such organisms are usually closely related Peripatric and...
, 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
Founder effect
In population genetics, the founder effect is the loss of genetic variation that occurs when a new population is established by a very small number of individuals from a larger population. It was first fully outlined by Ernst Mayr in 1942, using existing theoretical work by those such as Sewall...
causes rapid speciation after an increase in inbreeding
Inbreeding
Inbreeding is the reproduction from the mating of two genetically related parents. Inbreeding results in increased homozygosity, which can increase the chances of offspring being affected by recessive or deleterious traits. This generally leads to a decreased fitness of a population, which is...
increases selection on homozygotes, leading to rapid genetic change.
The third mechanism of speciation is parapatric speciation
Parapatric speciation
Parapatry is a term from biogeography, referring to organisms whose ranges do not significantly overlap but are immediately adjacent to each other; they only occur together in the narrow contact zone, if at all. This geographical distribution is opposed to sympatry & allopatry or peripatry...
. 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. Generally this occurs when there has been a drastic change in the environment within the parental species' habitat. One example is the grass Anthoxanthum odoratum
Anthoxanthum
Anthoxanthum, the vernal grasses or vernalgrasses, is a large genus of true grass with a cosmopolitan distribution.A. odoratum is a common species of acidic grassland and bogs in northern Europe....
, which can undergo parapatric speciation in response to localised metal pollution from mines. Here, plants evolve that have resistance to high levels of metals in the soil. 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
Character displacement
Character displacement refers to the phenomenon where differences among similar species whose distributions overlap geographically are accentuated in regions where the species co-occur but are minimized or lost where the species’ distributions do not overlap. This pattern results from evolutionary...
, which is when two species become more distinct in appearance.
Sympatric speciation
Sympatric speciation is the process through which new species evolve from a single ancestral species while inhabiting the same geographic region. In evolutionary biology and biogeography, sympatric and sympatry are terms referring to organisms whose ranges overlap or are even identical, so that...
species diverge without geographic isolation or changes in habitat. This form is rare since even a small amount of gene flow
Gene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...
may remove genetic differences between parts of a population. Generally, sympatric speciation in animals requires the evolution of both genetic differences
Polymorphism (biology)
Polymorphism in biology occurs when two or more clearly different phenotypes exist in the same population of a species — in other words, the occurrence of more than one form or morph...
and non-random mating
Assortative mating
Assortative mating , and the related concept Disassortative mating, is the phenomenon where a sexually reproducing organism chooses to mate with individuals that are similar or dissimilar to itself in some specific manner...
, to allow reproductive isolation to evolve.
One type of sympatric speciation involves cross-breeding 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
Meiosis
Meiosis is a special type of cell division necessary for sexual reproduction. The cells produced by meiosis are gametes or spores. The animals' gametes are called sperm and egg cells....
the homologous chromosome
Homologous chromosome
Homologous chromosomes are chromosome pairs of approximately the same length, centromere position, and staining pattern, with genes for the same characteristics at corresponding loci. One homologous chromosome is inherited from the organism's mother; the other from the organism's father...
s 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
Polyploidy
Polyploid is a term used to describe cells and organisms containing more than two paired sets of chromosomes. Most eukaryotic species are diploid, meaning they have two sets of chromosomes — one set inherited from each parent. However polyploidy is found in some organisms and is especially common...
. This allows the chromosomes from each parental species to form matching pairs during meiosis, since each parent's chromosomes are represented by a pair already. An example of such a speciation event is when the plant species Arabidopsis thaliana
Arabidopsis thaliana
Arabidopsis thaliana is a small flowering plant native to Europe, Asia, and northwestern Africa. A spring annual with a relatively short life cycle, arabidopsis is popular as a model organism in plant biology and genetics...
and Arabidopsis arenosa cross-bred to give the new species Arabidopsis suecica. This happened about 20,000 years ago, and the speciation process has been repeated in the laboratory, which allows the study of the genetic mechanisms involved in this process. Indeed, chromosome doubling within a species may be a common cause of reproductive isolation, as half the doubled chromosomes will be unmatched when breeding with undoubled organisms.
Speciation events are important in the theory of punctuated equilibrium
Punctuated equilibrium
Punctuated equilibrium is a theory in evolutionary biology which proposes that most species will exhibit little net evolutionary change for most of their geological history, remaining in an extended state called stasis...
, 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. In this theory, speciation and rapid evolution are linked, with natural selection and genetic drift acting most strongly on organisms undergoing speciation in novel habitats or small populations. 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
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...
is the disappearance of an entire species. Extinction is not an unusual event, as species regularly appear through speciation and disappear through extinction. Nearly all animal and plant species that have lived on Earth are now extinct, and extinction appears to be the ultimate fate of all species. These extinctions have happened continuously throughout the history of life, although the rate of extinction spikes in occasional mass extinction event
Extinction event
An extinction event is a sharp decrease in the diversity and abundance of macroscopic life. They occur when the rate of extinction increases with respect to the rate of speciation...
s. The Cretaceous–Tertiary extinction event, during which the non-avian dinosaurs went extinct, is the most well-known, but the earlier Permian–Triassic extinction event was even more severe, with approximately 96% of species driven to extinction. The Holocene extinction event
Holocene extinction event
The Holocene extinction refers to the extinction of species during the present Holocene epoch . The large number of extinctions span numerous families of plants and animals including mammals, birds, amphibians, reptiles and arthropods; a sizeable fraction of these extinctions are occurring in the...
is an ongoing mass extinction associated with humanity's expansion across the globe over the past few thousand years. Present-day extinction rates are 100–1000 times greater than the background rate and up to 30% of species may be extinct by the mid 21st century. Human activities are now the primary cause of the ongoing extinction event; global warming
Global warming
Global warming refers to the rising average temperature of Earth's atmosphere and oceans and its projected continuation. In the last 100 years, Earth's average surface temperature increased by about with about two thirds of the increase occurring over just the last three decades...
may further accelerate it in the future.
The role of extinction in evolution is not very well understood and may depend on which type of extinction is considered. The causes of the continuous "low-level" extinction events, which form the majority of extinctions, may be the result of competition between species for limited resources (competitive exclusion). If one species can out-compete another, this could produce species selection, with the fitter species surviving and the other species being driven to extinction. The intermittent mass extinctions are also important, but instead of acting as a selective force, they drastically reduce diversity in a nonspecific manner and promote bursts of rapid evolution
Adaptive radiation
In evolutionary biology, adaptive radiation is the evolution of ecological and phenotypic diversity within a rapidly multiplying lineage. Starting with a recent single ancestor, this process results in the speciation and phenotypic adaptation of an array of species exhibiting different...
and speciation in survivors.
Evolutionary history of life
Origin of life
Highly energetic chemistry is believed to have produced a self-replicating molecule around ago and half a billion years later the last common ancestor of all life existed. The current scientific consensusScientific consensus
Scientific consensus is the collective judgment, position, and opinion of the community of scientists in a particular field of study. Consensus implies general agreement, though not necessarily unanimity. Scientific consensus is not by itself a scientific argument, and it is not part of the...
is that the complex biochemistry
Biochemistry
Biochemistry, sometimes called biological chemistry, is the study of chemical processes in living organisms, including, but not limited to, living matter. Biochemistry governs all living organisms and living processes...
that makes up life came from simpler chemical reactions. The beginning of life may have included self-replicating molecules such as RNA
RNA
Ribonucleic acid , or RNA, is one of the three major macromolecules that are essential for all known forms of life....
, and the assembly of simple cells.
Common descent
All organismOrganism
In biology, an organism is any contiguous living system . In at least some form, all organisms are capable of response to stimuli, reproduction, growth and development, and maintenance of homoeostasis as a stable whole.An organism may either be unicellular or, as in the case of humans, comprise...
s on Earth
Earth
Earth is the third planet from the Sun, and the densest and fifth-largest of the eight planets in the Solar System. It is also the largest of the Solar System's four terrestrial planets...
are descended from a common ancestor or ancestral gene pool. Current species are a stage in the process of evolution, with their diversity the product of a long series of speciation and extinction events. The common descent
Common descent
In evolutionary biology, a group of organisms share common descent if they have a common ancestor. There is strong quantitative support for the theory that all living organisms on Earth are descended from a common ancestor....
of organisms was first deduced from four simple facts about organisms: 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. However, modern research has suggested that, due to horizontal gene transfer, this "tree of life
Tree of life (science)
Charles Darwin proposed that phylogeny, the evolutionary relatedness among species through time, was expressible as a metaphor he termed the Tree of Life...
" may be more complicated than a simple branching tree since some genes have spread independently between distantly related species.
Past species have also left records of their evolutionary history. Fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...
s, along with the comparative anatomy of present-day organisms, constitute the morphological, or anatomical, record. By comparing the anatomies of both modern and extinct species, paleontologists can infer the lineages of those species. 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
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
and archaea
Archaea
The Archaea are a group of single-celled microorganisms. A single individual or species from this domain is called an archaeon...
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
Biochemistry
Biochemistry, sometimes called biological chemistry, is the study of chemical processes in living organisms, including, but not limited to, living matter. Biochemistry governs all living organisms and living processes...
similarities between organisms. For example, all living cells use the same basic set of nucleotide
Nucleotide
Nucleotides are molecules that, when joined together, make up the structural units of RNA and DNA. In addition, nucleotides participate in cellular signaling , and are incorporated into important cofactors of enzymatic reactions...
s and amino acid
Amino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...
s. The development of molecular genetics
Molecular genetics
Molecular genetics is the field of biology and genetics that studies the structure and function of genes at a molecular level. The field studies how the genes are transferred from generation to generation. Molecular genetics employs the methods of genetics and molecular biology...
has revealed the record of evolution left in organisms' genome
Genome
In modern molecular biology and genetics, the genome is the entirety of an organism's hereditary information. It is encoded either in DNA or, for many types of virus, in RNA. The genome includes both the genes and the non-coding sequences of the DNA/RNA....
s: dating when species diverged through the molecular clock
Molecular clock
The molecular clock is a technique in molecular evolution that uses fossil constraints and rates of molecular change to deduce the time in geologic history when two species or other taxa diverged. It is used to estimate the time of occurrence of events called speciation or radiation...
produced by mutations. For example, these DNA sequence comparisons have revealed that humans and chimpanzees share 96% of their genomes and analyzing the few areas where they differ helps shed light on when the common ancestor of these species existed.
Evolution of life
ProkaryoteProkaryote
The prokaryotes are a group of organisms that lack a cell nucleus , or any other membrane-bound organelles. The organisms that have a cell nucleus are called eukaryotes. Most prokaryotes are unicellular, but a few such as myxobacteria have multicellular stages in their life cycles...
s inhabited the Earth from approximately 3–4 billion years ago. No obvious changes in morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
or cellular organisation occurred in these organisms over the next few billion years.
The eukaryotic cells
Eukaryote
A eukaryote is an organism whose cells contain complex structures enclosed within membranes. Eukaryotes may more formally be referred to as the taxon Eukarya or Eukaryota. The defining membrane-bound structure that sets eukaryotic cells apart from prokaryotic cells is the nucleus, or nuclear...
emerged between 1.6 – 2.7 billion years ago. The next major change in cell structure came when bacteria were engulfed by eukaryotic cells, in a cooperative association called endosymbiosis
Endosymbiont
An endosymbiont is any organism that lives within the body or cells of another organism, i.e. forming an endosymbiosis...
. The engulfed bacteria and the host cell then underwent co-evolution, with the bacteria evolving into either mitochondria
Mitochondrion
In cell biology, a mitochondrion is a membrane-enclosed organelle found in most eukaryotic cells. These organelles range from 0.5 to 1.0 micrometers in diameter...
or hydrogenosome
Hydrogenosome
A hydrogenosome is a membrane-enclosed organelle of some anaerobic ciliates, trichomonads and fungi. The hydrogenosomes of trichomonads produce molecular hydrogen, acetate, carbon dioxide and ATP by the combined actions of pyruvate:ferredoxin oxido-reductase, hydrogenase, acetate:succinate CoA...
s. Another engulfment of cyanobacterial-like organisms led to the formation of chloroplast
Chloroplast
Chloroplasts are organelles found in plant cells and other eukaryotic organisms that conduct photosynthesis. Chloroplasts capture light energy to conserve free energy in the form of ATP and reduce NADP to NADPH through a complex set of processes called photosynthesis.Chloroplasts are green...
s in algae and plants.
The history of life was that of the unicellular eukaryotes, prokaryotes and archaea until about 610 million years ago when multicellular organisms began to appear in the oceans in the Ediacaran
Ediacara biota
The Ediacara biota consisted of enigmatic tubular and frond-shaped, mostly sessile organisms which lived during the Ediacaran Period . Trace fossils of these organisms have been found worldwide, and represent the earliest known complex multicellular organisms.Simple multicellular organisms such as...
period. The evolution of multicellularity occurred in multiple independent events, in organisms as diverse as sponges, brown algae
Brown algae
The Phaeophyceae or brown algae , is a large group of mostly marine multicellular algae, including many seaweeds of colder Northern Hemisphere waters. They play an important role in marine environments, both as food and for the habitats they form...
, cyanobacteria, slime moulds and myxobacteria
Myxobacteria
The myxobacteria are a group of bacteria that predominantly live in the soil. The myxobacteria have very large genomes, relative to other bacteria, e.g. 9-10 million nucleotides. Sorangium cellulosum has the largest known bacterial genome, at 13.0 million nucleotides...
.
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
Cambrian explosion
The Cambrian explosion or Cambrian radiation was the relatively rapid appearance, around , of most major phyla, as demonstrated in the fossil record, accompanied by major diversification of other organisms, including animals, phytoplankton, and calcimicrobes...
. Here, the majority of types
Phylum
In biology, a phylum The term was coined by Georges Cuvier from Greek φῦλον phylon, "race, stock," related to φυλή phyle, "tribe, clan." is a taxonomic rank below kingdom and above class. "Phylum" is equivalent to the botanical term division....
of modern animals appeared in the fossil record, as well as unique lineages that subsequently became extinct. Various triggers for the Cambrian explosion have been proposed, including the accumulation of oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...
in the atmosphere
Atmosphere
An atmosphere is a layer of gases that may surround a material body of sufficient mass, and that is held in place by the gravity of the body. An atmosphere may be retained for a longer duration, if the gravity is high and the atmosphere's temperature is low...
from photosynthesis
Photosynthesis
Photosynthesis is a chemical process that converts carbon dioxide into organic compounds, especially sugars, using the energy from sunlight. Photosynthesis occurs in plants, algae, and many species of bacteria, but not in archaea. Photosynthetic organisms are called photoautotrophs, since they can...
.
About 500 million years ago, plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
s and fungi
Fungus
A fungus is a member of a large group of eukaryotic organisms that includes microorganisms such as yeasts and molds , as well as the more familiar mushrooms. These organisms are classified as a kingdom, Fungi, which is separate from plants, animals, and bacteria...
colonised the land and were soon followed by arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...
s and other animals. Insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s were particularly successful and even today make up the majority of animal species. Amphibian
Amphibian
Amphibians , are a class of vertebrate animals including animals such as toads, frogs, caecilians, and salamanders. They are characterized as non-amniote ectothermic tetrapods...
s first appeared around 364 million years ago, followed by early amniote
Amniote
The amniotes are a group of tetrapods that have a terrestrially adapted egg. They include synapsids and sauropsids , as well as their fossil ancestors. Amniote embryos, whether laid as eggs or carried by the female, are protected and aided by several extensive membranes...
s, then bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...
s around 155 million years ago (both from "reptile
Reptile
Reptiles are members of a class of air-breathing, ectothermic vertebrates which are characterized by laying shelled eggs , and having skin covered in scales and/or scutes. They are tetrapods, either having four limbs or being descended from four-limbed ancestors...
"-like lineages), mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...
s around 129 million years ago, homininae
Homininae
Homininae is a subfamily of Hominidae, which includes humans, gorillas and chimpanzees, and some extinct relatives; it comprises all those hominids, such as Australopithecus, that arose after the split from orangutans . Our family tree, which has 3 main branches leading to chimpanzees, humans and...
around 10 million years ago and modern humans around 0.25 million years ago. 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
Biomass (ecology)
Biomass, in ecology, is the mass of living biological organisms in a given area or ecosystem at a given time. Biomass can refer to species biomass, which is the mass of one or more species, or to community biomass, which is the mass of all species in the community. It can include microorganisms,...
and species being prokaryotes.
Applications
Concepts and models used in evolutionary biology, in particular natural selection, have many applications.Artificial selection
Artificial selection
Artificial selection describes intentional breeding for certain traits, or combination of traits. The term was utilized by Charles Darwin in contrast to natural selection, in which the differential reproduction of organisms with certain traits is attributed to improved survival or reproductive...
is the intentional selection of traits in a population of organisms. This has been used for thousands of years in the domestication
Domestication
Domestication or taming is the process whereby a population of animals or plants, through a process of selection, becomes accustomed to human provision and control. In the Convention on Biological Diversity a domesticated species is defined as a 'species in which the evolutionary process has been...
of plants and animals. More recently, such selection has become a vital part of genetic engineering
Genetic engineering
Genetic engineering, also called genetic modification, is the direct human manipulation of an organism's genome using modern DNA technology. It involves the introduction of foreign DNA or synthetic genes into the organism of interest...
, with selectable marker
Selectable marker
A selectable marker is a gene introduced into a cell, especially a bacterium or to cells in culture, that confers a trait suitable for artificial selection. They are a type of reporter gene used in laboratory microbiology, molecular biology, and genetic engineering to indicate the success of a...
s such as antibiotic resistance genes being used to manipulate DNA. In repeated rounds of mutation and selection proteins with valuable properties have evolved, for example modified enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
s and new antibodies
Antibody
An antibody, also known as an immunoglobulin, is a large Y-shaped protein used by the immune system to identify and neutralize foreign objects such as bacteria and viruses. The antibody recognizes a unique part of the foreign target, termed an antigen...
, in a process called directed evolution
Directed evolution
thumb|250px|right|An example of a possible round to evolve a protein based fluorescent sensor for a specific analyte using two consecutive FACS sortings...
.
Understanding the changes that have occurred during organism's evolution can reveal the genes needed to construct parts of the body, genes which may be involved in human genetic disorder
Genetic disorder
A genetic disorder is an illness caused by abnormalities in genes or chromosomes, especially a condition that is present from before birth. Most genetic disorders are quite rare and affect one person in every several thousands or millions....
s. For example, the mexican tetra
Mexican tetra
The Mexican tetra or Blind Cave Fish is a freshwater fishof the characin family oforder Characiformes....
is an albino cavefish that lost its eyesight during evolution. 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. This helped identify genes required for vision and pigmentation.
In computer science
Computer science
Computer science or computing science is the study of the theoretical foundations of information and computation and of practical techniques for their implementation and application in computer systems...
, simulations of evolution using evolutionary algorithm
Evolutionary algorithm
In artificial intelligence, an evolutionary algorithm is a subset of evolutionary computation, a generic population-based metaheuristic optimization algorithm. An EA uses some mechanisms inspired by biological evolution: reproduction, mutation, recombination, and selection...
s and artificial life
Artificial life
Artificial life is a field of study and an associated art form which examine systems related to life, its processes, and its evolution through simulations using computer models, robotics, and biochemistry. The discipline was named by Christopher Langton, an American computer scientist, in 1986...
started in the 1960s and was extended with simulation of artificial selection
Artificial selection
Artificial selection describes intentional breeding for certain traits, or combination of traits. The term was utilized by Charles Darwin in contrast to natural selection, in which the differential reproduction of organisms with certain traits is attributed to improved survival or reproductive...
. Artificial evolution
Evolutionary algorithm
In artificial intelligence, an evolutionary algorithm is a subset of evolutionary computation, a generic population-based metaheuristic optimization algorithm. An EA uses some mechanisms inspired by biological evolution: reproduction, mutation, recombination, and selection...
became a widely recognised optimisation method as a result of the work of Ingo Rechenberg
Ingo Rechenberg
Ingo Rechenberg is a German computer scientist and professor. Rechenberg is a pioneer of the fields of evolutionary computation and artificial evolution. In the 1960s and 1970s he invented a highly influential set of optimization methods known as evolution strategies...
in the 1960s. He used evolution strategies
Evolution strategy
In computer science, evolution strategy is an optimization technique based on ideas of adaptation and evolution. It belongs to the general class of evolutionary computation or artificial evolution methodologies.-History:...
to solve complex engineering problems. Genetic algorithm
Genetic algorithm
A genetic algorithm is a search heuristic that mimics the process of natural evolution. This heuristic is routinely used to generate useful solutions to optimization and search problems...
s in particular became popular through the writing of John Holland
John Henry Holland
John Henry Holland is an American scientist and Professor of Psychology and Professor of Electrical Engineering and Computer Science at the University of Michigan, Ann Arbor. He is a pioneer in complex systems and nonlinear science. He is known as the father of genetic algorithms. He was awarded...
. Practical applications also include automatic evolution of computer programs
Genetic programming
In artificial intelligence, genetic programming is an evolutionary algorithm-based methodology inspired by biological evolution to find computer programs that perform a user-defined task. It is a specialization of genetic algorithms where each individual is a computer program...
. Evolutionary algorithms are now used to solve multi-dimensional problems more efficiently than software produced by human designers and also to optimise the design of systems.
Social and cultural responses
In the 19th century, particularly after the publication of On the Origin of Species in 1859, the idea that life had evolved was an active source of academic debate centred on the philosophical, social and religious implications of evolution. Nowadays, the modern evolutionary synthesis is accepted by a vast majority of scientists. However, evolution remains a contentious concept for some theistsTheism
Theism, in the broadest sense, is the belief that at least one deity exists.In a more specific sense, theism refers to a doctrine concerning the nature of a monotheistic God and God's relationship to the universe....
.
While various religions and denominations have reconciled their beliefs with evolution through concepts such as theistic evolution
Theistic evolution
Theistic evolution or evolutionary creation is a concept that asserts that classical religious teachings about God are compatible with the modern scientific understanding about biological evolution...
, there are creationists
Creationism
Creationism is the religious beliefthat humanity, life, the Earth, and the universe are the creation of a supernatural being, most often referring to the Abrahamic god. As science developed from the 18th century onwards, various views developed which aimed to reconcile science with the Genesis...
who believe that evolution is contradicted by the creation myths found in their religion
Religion
Religion is a collection of cultural systems, belief systems, and worldviews that establishes symbols that relate humanity to spirituality and, sometimes, to moral values. Many religions have narratives, symbols, traditions and sacred histories that are intended to give meaning to life or to...
s and who raise various objections to evolution
Objections to evolution
Objections to evolution have been raised since evolutionary ideas came to prominence in the 19th century. When Charles Darwin published his 1859 book On the Origin of Species, his theory of evolution by natural selection initially met opposition from scientists with different theories, but came to...
. As had been demonstrated by responses to the publication of Vestiges of the Natural History of Creation
Vestiges of the Natural History of Creation
Vestiges of the Natural History of Creation is a unique work of speculative natural history published anonymously in England in 1844. It brought together various ideas of stellar evolution with the progressive transmutation of species in an accessible narrative which tied together numerous...
in 1844, the most controversial aspect of evolutionary biology is the implication of human evolution
Human evolution
Human evolution refers to the evolutionary history of the genus Homo, including the emergence of Homo sapiens as a distinct species and as a unique category of hominids and mammals...
that humans share common ancestry with apes and that the mental and moral faculties of humanity have the same types of natural causes as other inherited traits in animals. In some countries, notably the United States, these tensions between science and religion have fuelled the current creation-evolution controversy, a religious conflict focusing on politics
Politics of creationism
Different churches address the theological implications raised by creationism and evolution in different ways.-Evolution:Most contemporary Christian leaders and scholars from many mainstream churches, such as Roman Catholic, Anglican and some Lutheran denominations, reject reading the Bible as...
and public education
Creation and evolution in public education
The status of creation and evolution in public education has been the subject of substantial debate in legal, political, and religious circles...
. While other scientific fields such as cosmology
Physical cosmology
Physical cosmology, as a branch of astronomy, is the study of the largest-scale structures and dynamics of the universe and is concerned with fundamental questions about its formation and evolution. For most of human history, it was a branch of metaphysics and religion...
and Earth science
Earth science
Earth science is an all-embracing term for the sciences related to the planet Earth. It is arguably a special case in planetary science, the Earth being the only known life-bearing planet. There are both reductionist and holistic approaches to Earth sciences...
also conflict with literal interpretations of many religious texts, evolutionary biology experiences significantly more opposition from religious literalists.
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
Scopes Trial
The Scopes Trial—formally known as The State of Tennessee v. John Thomas Scopes and informally known as the Scopes Monkey Trial—was a landmark American legal case in 1925 in which high school science teacher, John Scopes, was accused of violating Tennessee's Butler Act which made it unlawful to...
decision of 1925 caused the subject to become very rare in American secondary biology textbooks for a generation, but it was gradually re-introduced about a generation later and legally protected with the 1968 Epperson v. Arkansas
Epperson v. Arkansas
Epperson v. Arkansas, 393 U.S. 97 , was a United States Supreme Court case that invalidated an Arkansas statute that prohibited the teaching of human evolution in the public schools...
decision. Since then, the competing religious belief of creationism was legally disallowed in secondary school curricula in various decisions in the 1970s and 1980s, but it returned in the form of intelligent design
Intelligent design
Intelligent design is the proposition that "certain features of the universe and of living things are best explained by an intelligent cause, not an undirected process such as natural selection." It is a form of creationism and a contemporary adaptation of the traditional teleological argument for...
, to be excluded once again in the 2005 Kitzmiller v. Dover Area School District
Kitzmiller v. Dover Area School District
Tammy Kitzmiller, et al. v. Dover Area School District, et al. was the first direct challenge brought in the United States federal courts testing a public school district policy that required the teaching of intelligent design...
case.
See also
- Applications of evolutionApplications of evolutionEvolutionary biology, in particular the understanding of how organisms evolve through natural selection, is an area of science with many practical applications.-Wider biology:...
- Biocultural evolutionBiocultural evolutionBiocultural evolution refers to historical evolutionary processes that occur as a result of culture's interaction with biology.-Cultural factors:...
- Biological imperative
- Current research in evolutionary biologyCurrent research in evolutionary biologyCurrent research in evolutionary biology covers diverse topics, as should be expected given the centrality of evolution to understanding biology...
- Evolutionary anthropologyEvolutionary anthropologyEvolutionary anthropology is the interdisciplinary study of evolution of human physiology and human behaviour and the relation between hominids and non-hominid primates. Evolutionary anthropology is based in natural science and social science...
- Evolutionary neuroscienceEvolutionary neuroscienceEvolutionary neuroscience is an interdisciplinary scientific research field that studies the evolution of nervous systems. Evolutionary neuroscientists attempt to understand the evolution and natural history of nervous system structure and function. The field draws on concepts and findings from...
- Evolutionary psychologyEvolutionary psychologyEvolutionary psychology is an approach in the social and natural sciences that examines psychological traits such as memory, perception, and language from a modern evolutionary perspective. It seeks to identify which human psychological traits are evolved adaptations, that is, the functional...
- Evolutionary history of lifeEvolutionary history of lifeThe evolutionary history of life on Earth traces the processes by which living and fossil organisms have evolved since life on Earth first originated until the present day. Earth formed about 4.5 Ga and life appeared on its surface within one billion years...
- Human evolutionHuman evolutionHuman evolution refers to the evolutionary history of the genus Homo, including the emergence of Homo sapiens as a distinct species and as a unique category of hominids and mammals...
- NeurocultureNeurocultureNeuroCulture is the relation between the sciences that study the functioning of the brain and culture. We understanding the latter as the knowledge, history, habits, ideas and values of the human race, and their manifestations in any form of expression: social, scientific, artistic,...
- SociobiologySociobiologySociobiology is a field of scientific study which is based on the assumption that social behavior has resulted from evolution and attempts to explain and examine social behavior within that context. Often considered a branch of biology and sociology, it also draws from ethology, anthropology,...
- Sociocultural evolutionSociocultural evolutionSociocultural evolution is an umbrella term for theories of cultural evolution and social evolution, describing how cultures and societies have changed over time...
- Technological evolutionTechnological evolutionTechnological evolution is the name of a science and technology studies theory describing technology development, developed by Czech philosopher Radovan Richta.-Theory of technological evolution:...