Everything about Theory Of Evolution totally explained
In
biology,
evolution is the process of change in the
inherited traits of a
population of organisms from one
generation to the next. The
genes that are passed on to an organism's offspring
produce the inherited traits that are the basis of evolution.
Mutations in genes can produce new or altered traits in individuals, resulting in the appearance of
heritable differences between organisms, but new traits also come from the transfer of genes between populations, as in
migration, or between species, in
horizontal gene transfer. In species that reproduce
sexually, new combinations of genes are produced by
genetic recombination, which can increase the variation in traits between organisms. Evolution occurs when these heritable differences become more common or rare in a population.
There are two major mechanisms driving evolution. The first is
natural selection, which is a process causing heritable traits that are helpful for survival and reproduction to become more common in a population, and harmful traits to become more rare. This occurs because individuals with advantageous traits are more likely to reproduce successfully, so that more individuals in the next generation inherit these traits. Over many generations,
adaptations occur through a combination of successive, small, random changes in traits, and natural selection of those variants best-suited for their environment. In contrast,
genetic drift produces random changes in the frequency of traits in a population. Genetic drift results from the role chance plays in whether a given individual will survive and reproduce. Though the changes produced in any one generation by drift and selection are small, differences accumulate with each subsequent generation and can, over time, cause substantial changes in the organisms.
One definition of a
species is a group of organisms that can reproduce with one another and produce fertile offspring. When a species is separated into populations that are
prevented from interbreeding, mutations, genetic drift, and natural selection cause the accumulation of differences over generations and the
emergence of new species. The similarities between organisms suggest that all known species are
descended from a common ancestor (or ancestral gene pool) through this process of gradual divergence.
Evolutionary biology documents the fact that evolution occurs, and also develops and tests
theories that explain why it occurs. Studies of the
fossil record and the
diversity of living organisms had convinced most scientists by the mid-nineteenth century that species changed over time. However, the mechanism driving these changes remained unclear until the 1859 publication of
Charles Darwin's
On the Origin of Species, detailing the
theory of evolution by natural selection. Darwin's work soon led to overwhelming acceptance of evolution within the scientific community. In the 1930s, Darwinian natural selection was combined with
Mendelian inheritance to form the
modern evolutionary synthesis,
Heredity
Inheritance in organisms occurs through discrete
traits – particular characteristics of an organism. In humans, for example,
eye color is an inherited characteristic, which individuals can inherit from one of their parents. Inherited traits are controlled by
genes 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 behavior of an organism is called its
phenotype. These traits come from the interaction of its genotype with the environment. As a result, not every aspect of an organism's phenotype is inherited.
Suntanned skin results from the interaction between a person's genotype and sunlight; thus, suntans are not passed on to people's children. However, people have different responses to sunlight, arising from differences in their genotype; a striking example is individuals with the inherited trait of
albinism, who don't tan and are highly sensitive to
sunburn.
Heritable traits are propagated between generations via
DNA, a
molecule which is capable of encoding genetic information.
Variation
Because an individual's
phenotype results from the interaction of its
genotype with the environment, the variation in phenotypes in a population reflects the variation in these organisms' genotypes.
Variation comes from
mutations in
genetic material, migration between populations (
gene flow), and the reshuffling of genes through
sexual reproduction. Variation also comes from exchanges of genes between different species; for example, through
horizontal gene transfer in
bacteria, and
hybridization in plants. Despite the constant introduction of variation through these processes, most of the
genome of a species is identical in all individuals of that species. However, even relatively small changes in genotype can lead to dramatic changes in phenotype: chimpanzees and humans differ in only about 5% of their genomes.
Mutation
Genetic variation comes from
random mutations that occur in the genomes of organisms. Mutations are changes in the DNA sequence of a cell's genome and are caused by
radiation,
viruses,
transposons and
mutagenic chemicals, as well as errors that occur during
meiosis or
DNA replication. These mutagens produce several different types of change in DNA sequences; these can either have no effect, alter the
product of a gene, or prevent the gene from functioning. Studies in the fly
Drosophila melanogaster suggest that if a mutation changes a protein produced by a gene, this will probably be harmful, with about 70 percent of these mutations having damaging effects, and the remainder being either neutral or weakly beneficial. Due to the damaging effects that mutations can have on cells, organisms have evolved mechanisms such as
DNA repair to remove mutations. Some species such as
retroviruses have such high mutation rates that most of their offspring will possess a mutated gene. Such rapid mutation may have been selected so that these viruses can constantly and rapidly evolve, and thus evade the responses of the human
immune system.
Mutations can involve large sections of DNA becoming
duplicated, which is a major source of raw material for evolving new genes, with tens to hundreds of genes duplicated in animal genomes every million years. Most genes belong to larger
families of genes of
shared ancestry. Novel genes are produced either through duplication and mutation of an ancestral gene, or by recombining parts of different genes to form new combinations with new functions. For example, the human eye uses four genes to make structures that sense light: three for
color vision and one for
night vision; all four arose from a single ancestral gene. An advantage of duplicating a gene (or even an
entire genome) is that overlapping or
redundant functions in multiple genes allows alleles to be retained that would otherwise be harmful, thus increasing genetic diversity.
Changes in chromosome number may involve even larger mutations, where long segments of the DNA within chromosomes breaks and then rearranges. For example, two chromosomes in the
Homo genus fused to produce human
chromosome 2; this fusion didn't occur in the
lineage of the other apes, and they retain these separate chromosomes. In evolution, the most important role of such chromosomal rearrangements may be to accelerate the divergence of a population into new species by making populations less likely to interbreed, and thereby preserving genetic differences between these populations.
Sequences of DNA that can move about the genome, such as
transposons, make up a major fraction of the genetic material of plants and animals, and may have been important in the evolution of genomes. For example, more than a million copies of the
Alu sequence are present in the
human genome, and these sequences have now been recruited to perform functions such as regulating
gene expression. Another effect of these mobile DNA sequences is that when they move within a genome, they can mutate or delete existing genes and thereby produce genetic diversity.
Sex and recombination
In asexual organisms, genes are inherited together, or
linked, as they can't mix with genes in other organisms during reproduction. However, the offspring of
sexual organisms contain random mixtures of their parents' chromosomes that are produced through
independent assortment. In the related process of
genetic recombination, sexual organisms can also exchange DNA between two matching chromosomes. Recombination and reassortment don't alter allele frequencies, but instead change which alleles are associated with each other, producing offspring with new combinations of alleles. While this process increases the variation in any individual's offspring, genetic mixing can be predicted to either have no effect, increase, or decrease the
genetic variation in the population, depending on how the various alleles in the population are distributed. For example, if two alleles are randomly distributed in a population, then sex will have no effect on variation; however, if two alleles tend to be found as a pair, then genetic mixing will even out this non-random distribution and over time make the organisms in the population more similar to each other.
Recombination allows even alleles that are close together in a strand of DNA to be
inherited independently. However, the rate of recombination is low, since in humans in a stretch of DNA one million
base pairs long there's about a one in a hundred chance of a recombination event occurring 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. This tendency is measured by finding how often two alleles occur together, which is called their
linkage disequilibrium. A set of alleles that's usually inherited in a group is called a
haplotype.
Sexual reproduction helps to remove harmful mutations and retain beneficial mutations. Consequently, when alleles can't be separated by recombination – such as in mammalian
Y chromosomes, which pass intact from fathers to sons – harmful
mutations accumulate. In addition, recombination and reassortment can produce individuals with new and advantageous gene combinations. These positive effects are balanced by the fact that this process can cause mutations and separate beneficial combinations of genes. A
population is a localized group of individuals belonging to the same species. For example, all of the moths of the same species living in an isolated forest represent a population. A single gene in this population may have several alternate forms, which account for variations between the phenotypes of the organisms. An example might be a gene for coloration in moths that has two alleles: black and white. A
gene pool is the complete set of alleles in a single population, so each allele occurs a certain number of times in a gene pool. The fraction of genes within the gene pool that are a particular allele is called the
allele frequency. Evolution occurs when there are changes in the frequencies of alleles within a population of interbreeding organisms; for example the allele for black color in a population of moths becoming more common.
To understand the mechanisms that cause a population to evolve, it's useful to consider what conditions are required for a population not to evolve. The
Hardy-Weinberg principle states that the frequencies of alleles (variations in a gene) in a sufficiently large population will remain constant if the only forces acting on that population are the random reshuffling of alleles during the formation of the sperm or egg, and the random combination of the alleles in these sex cells during
fertilization. Such a population is said to be in
Hardy-Weinberg equilibrium - it isn't evolving.
Mechanisms
There are three basic mechanisms of evolutionary change:
natural selection,
genetic drift, and
gene flow. Natural selection favors genes that improve capacity for survival and reproduction. Genetic drift is random change in the frequency of alleles, caused by the random sampling of a generation's genes during reproduction, and gene flow is the transfer of genes within and between populations. The relative importance of natural selection and genetic drift in a population varies depending on the strength of the selection and the
effective population size, which is the number of individuals capable of breeding. Natural selection usually predominates in large populations, while genetic drift dominates in small populations. The dominance of genetic drift in small populations can even lead to the fixation of slightly deleterious mutations. As a result, changing population size can dramatically influence the course of evolution.
Population bottlenecks, where the population shrinks temporarily and therefore loses genetic variation, result in a more uniform population. Consequently, 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're "selected
against". 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's selection against extreme trait values on both ends, which causes a decrease in
variance around the average value. 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 colors 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.
An active area of research is the
unit of selection, with natural selection being proposed to work at the level of genes, cells, individual organisms, groups of organisms and even species. None of these models are mutually-exclusive and selection may act on multiple levels simultaneously. Below the level of the individual, genes called transposons try to copy themselves throughout the
genome. Selection at a level above the individual, such as
group selection, may allow the evolution of co-operation, as discussed below.
Genetic drift
Genetic drift is the change in allele frequency from one generation to the next that occurs because alleles in offspring are a
random sample of those in the parents, as well as from the role that chance plays in determining whether a given individual will survive and reproduce.
The time for an allele to become fixed by genetic drift depends on population size, with fixation occurring more rapidly in smaller populations. The precise measure of populations that's important here's called the
effective population size, which was defined by
Sewall Wright as a theoretical number representing the number of breeding individuals that would exhibit the same observed degree of inbreeding.
Although natural selection is responsible for adaptation, the relative importance of the two forces of natural selection and genetic drift in driving evolutionary change in general is an area of current research in evolutionary biology. These investigations were prompted by the
neutral theory of molecular evolution, which proposed that most evolutionary changes are the result of the fixation of
neutral mutations that don't have any immediate effects on the fitness of an organism. Hence, in this model, most genetic changes in a population are the result of constant mutation pressure and genetic drift.
Gene flow
Gene flow is the exchange of genes between populations, which are usually of the same species. Examples of gene flow within a species include the migration and then breeding of organisms, or the exchange of
pollen. Gene transfer between species includes the formation of
hybrid organisms and
horizontal gene transfer.
Migration into or out of a population can change allele frequencies, as well as introducing genetic variation into a population. Immigration may add new genetic material to the established
gene pool of a population. Conversely, emigration may remove genetic material. As
barriers to reproduction between two diverging populations are required for the populations to
become new species, gene flow may slow this process by spreading genetic differences between the populations. Gene flow is hindered by mountain ranges, oceans and deserts or even man-made structures such as the
Great Wall of China, which has hindered the flow of plant genes.
Depending on how far two species have diverged since their
most recent common ancestor, it may still be possible for them to produce offspring, as with
horses and
donkeys mating to produce
mules. Such
hybrids are generally
infertile, due to the two different sets of chromosomes being unable to pair up during
meiosis. 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 hybridization in creating
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.
Hybridization is, however, an important means of speciation in plants, since
polyploidy (having more than two copies of each chromosome) is tolerated in plants more readily than in animals. Polyploidy is important in hybrids as it allows reproduction, with the two different sets of chromosomes each being able to pair with an identical partner during meiosis. Polyploids also have more genetic diversity, which allows them to avoid
inbreeding depression in small populations.
Horizontal gene transfer is the transfer of genetic material from one organism to another organism that isn't 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 may also have occurred.
Viruses can also carry DNA between organisms, allowing transfer of genes even across
biological domains. Gene transfer has also occurred between the ancestors of
eukaryotic cells and prokaryotes, during the acquisition of the
chloroplast and
mitochondrial.
Outcomes
Evolution influences every aspect of the form and behavior of organisms. Most prominent are the specific behavioral and physical
adaptations that are the outcome of natural selection. These adaptations increase fitness by aiding activities such as finding food, avoiding predators or attracting mates. Organisms can also respond to selection by
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 can't or won't interbreed.
These outcomes of evolution are sometimes divided into
macroevolution, which is evolution that occurs at or above the level of species, such as
speciation, and
microevolution, which is smaller evolutionary changes, such as adaptations, within a species or population. In general, macroevolution is the outcome of long periods of microevolution. Thus, the distinction between micro- and macroevolution isn't a fundamental one - the difference is simply the time involved. However, in macroevolution, the traits of the entire species are 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 can sometimes be separate.
A common misconception is that evolution is "progressive," but natural selection has no long-term goal and doesn't necessarily produce greater complexity. Although
complex species have evolved, this occurs as a side effect of the overall number of organisms increasing, and simple forms of life remain more common. For example, the overwhelming majority of species are microscopic
prokaryotes, 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's
more noticeable.
Adaptation
Adaptations are structures or behaviors that enhance a specific function, causing organisms to become better at surviving and reproducing. This process can 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 mutations causing
antibiotic resistance by either modifying the target of the drug, or removing the transporters that allow the drug into the cell. However, many traits that appear to be simple adaptations are in fact
exaptations: 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
Holapsis 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.]]
As adaptation occurs through the gradual modification of existing structures, structures with similar internal organization may have very 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 structurally similar to both human hands and seal flippers, due to the common descent of these structures from an ancestor that also had five digits at the end of each forelimb. Other idiosyncratic anatomical features, such as
bones in the wrist of the
panda being formed into a false "thumb," indicate that an organism's evolutionary lineage can limit what adaptations are possible.
During adaptation, some structures may lose their original function and become
vestigial structures. 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 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, 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 create 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
chickens causing embryos to grow teeth similar to those of
crocodiles.
Co-evolution
Interactions between organisms can produce both conflict and co-operation. When the interaction is between pairs of species, such as a
pathogen and a
host, or a
predator and its prey, these species can develop matched sets of adaptations. Here, the evolution of one species causes adaptations in a second species. These changes in the second species then, in turn, cause new adaptations in the first species. This cycle of selection and response is called
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 resistance in the snake.
Co-operation
However, not all interactions between species involve conflict. Many cases of mutually beneficial interactions have evolved. For instance, an extreme cooperation exists between plants and the
mycorrhizal fungi that grow on their roots and aid the plant in absorbing nutrients from the soil. 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
bees,
termites and
ants, where sterile insects feed and guard the small number of organisms in a
colony that are able to reproduce. On an even smaller scale, the
somatic cells that make up the body of an animal are limited in their capacity to reproduce in order to maintain a stable organism, which then supports a small number of the animal's
germ cells to produce offspring. Here, somatic cells respond to specific signals that instruct them to either
grow or
kill themselves. If cells ignore these signals and attempt to multiply inappropriately, their uncontrolled growth causes
cancer. This activity is selected for because if the
helping individual contains alleles which promote the helping activity, it's 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.
Speciation
Speciation is the process where a species diverges into two or more descendant species. It 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. As selection and drift act independently in isolated populations, separation will eventually produce organisms that can't 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 through both rapid genetic drift and selection on a small gene pool.
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's no physical separation between these two populations. Instead, speciation results from the evolution of mechanisms that reduce gene flow between the two populations. Here, plants evolve that have resistance to high levels of metals in the soil. Selection against interbreeding with the metal-sensitive parental population produces a change in flowering time of the metal-resistant plants, causing 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 isn't common in animals as animal hybrids are usually sterile, because during
meiosis the
homologous chromosomes from each parent, being from different species can't successfully pair. It is more common in plants, however because plants often double their number of chromosomes, to form
polyploids. This allows the chromosomes from each parental species to form a matching pair during meiosis, as each parent's chromosomes is 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. 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 is the disappearance of an entire species. Extinction isn't an unusual event, as species regularly appear through speciation, and disappear through extinction. Indeed, virtually all animal and plant species that have lived on earth are now extinct. These extinctions have happened continuously throughout the history of life, although the rate of extinction spikes in occasional mass
extinction events. The
Cretaceous–Tertiary extinction event, during which the dinosaurs went extinct, is the most well-known, but the earlier
Permian–Triassic extinction event was even more severe, with approximately 96 percent of species driven to extinction. 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 depends on which type is considered. The causes of the continuous "low-level" extinction events, which form the majority of extinctions, are not well understood and may be the result of competition between species for shared resources. The current
scientific consensus is that the complex
biochemistry that makes up life came from simpler chemical reactions, but it's unclear how this occurred. Not much is certain about the earliest developments in life, the structure of the first living things, or the identity and nature of any
last universal common ancestor or ancestral gene pool. Consequently, there's no scientific consensus on how life began, but proposals include self-replicating molecules such as
RNA, and the assembly of simple cells.
Common descent
All
organisms 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've geographic distributions that can't be explained by local adaptation. Second, the diversity of life isn't 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. 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 don't 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
nucleic acids and
amino acids. The development of
molecular genetics has revealed the record of evolution left in organisms'
genomes: dating when species diverged through the
molecular clock produced by mutations. For example, these DNA sequence comparisons have revealed the close genetic similarity between humans and chimpanzees and shed light on when the common ancestor of these species existed.
Evolution of life
Despite the uncertainty on how life began, it's clear that
prokaryotes were the first organisms to inhabit Earth, approximately 3–4 billion years ago. No obvious changes in
morphology or cellular organization occurred in these organisms over the next few billion years.
The
eukaryotes were the next major innovation in evolution. These came from ancient bacteria being engulfed by the ancestors of eukaryotic cells, in a cooperative association called
endosymbiosis. The engulfed bacteria and the host cell then underwent co-evolution, with the bacteria evolving into either
mitochondria or
hydrogenosomes. An independent second engulfment of
cyanobacterial-like organisms led to the formation of
chloroplasts in algae and plants.
The history of life was that of the unicellular eukaryotes, prokaryotes, and archaea until about a billion years ago when multicellular organisms began to appear in the oceans in the
Ediacaran period. 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,
plants and
fungi colonized the land, and were soon followed by
arthropods and other animals.
Amphibians first appeared around 300 million years ago, followed by early
amniotes, then
mammals around 200 million years ago and
birds around 100 million years ago (both from "
reptile"-like lineages). However, despite the evolution of these large animals, smaller organisms similar to the types that evolved early in this process continue to be highly successful and dominate the Earth, with the majority of both
biomass and species being prokaryotes. Others who considered such ideas included the Greek philosopher
Empedocles, the
Roman philosopher-poet Lucretius, the
Arab biologist Al-Jahiz, the
Persian philosopher Ibn Miskawayh, the
Brethren of Purity, and the Eastern philosopher
Zhuangzi. As biological knowledge grew in the 18th century, evolutionary ideas were set out by a few natural philosophers including
Pierre Maupertuis in 1745 and
Erasmus Darwin in 1796. The ideas of the biologist
Jean-Baptiste Lamarck about
transmutation of species had wide influence.
Charles Darwin formulated his idea of
natural selection in 1838 and was still developing his theory in 1858 when
Alfred Russel Wallace sent him a similar theory, and both were presented to the
Linnean Society of London in
separate papers. At the end of 1859 Darwin's publication of
On the Origin of Species explained natural selection in detail and presented evidence leading to increasingly wide acceptance of the occurrence of evolution.
Debate about the mechanisms of evolution continued, and Darwin couldn't explain the source of the heritable variations which would be acted on by natural selection. Like Lamarck, he thought that parents
passed on adaptations acquired during their lifetimes, a theory which was subsequently dubbed
Lamarckism. In the 1880s
August Weismann's experiments indicated that changes from use and disuse were not heritable, and Lamarckism gradually fell from favour. More significantly, Darwin couldn't account for how traits were passed down from generation to generation. In 1865
Gregor Mendel found that traits were
inherited in a predictable manner. When Mendel's work was rediscovered in 1900, disagreements over the rate of evolution predicted by early geneticists and
biometricians led to a rift between the Mendelian and Darwinian models of evolution.
This contradiction was reconciled in the 1930s by biologists such as
Ronald Fisher. The end result was a combination of evolution by natural selection and Mendelian inheritance, the
modern evolutionary synthesis. In the 1940s, the identification of
DNA as the genetic material by
Oswald Avery and colleagues and the subsequent publication of the structure of DNA by
James Watson and
Francis Crick in 1953, demonstrated the physical basis for inheritance. Since then,
genetics and
molecular biology have become core parts of
evolutionary biology and have revolutionized the field of
phylogenetics.
In its early history, evolutionary biology primarily drew in scientists from traditional taxonomically-oriented disciplines, whose specialist training in particular organisms addressed general questions in evolution. As evolutionary biology expanded as an academic discipline, particularly after the development of the modern evolutionary synthesis, it began to draw more widely from the biological sciences.
Although
many religions and denominations have reconciled their beliefs with evolution through various concepts of
theistic evolution, there are many
creationists who believe that evolution is contradicted by the
creation myths found in their respective religions. As Darwin recognized early on, the most controversial aspect of evolutionary thought is its implications for
human origins. In some countries—notably the United States—these tensions between scientific and religious teachings have fueled the ongoing
creation–evolution controversy, a religious conflict focusing on
politics and
public education. 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 many religious believers.
Evolution has been used to support philosophical positions that promote
discrimination and
racism. For example, the
eugenic ideas of
Francis Galton were developed to argue that the human gene pool should be improved by
selective breeding policies, including incentives for those considered "good stock" to reproduce, and the
compulsory sterilization,
prenatal testing,
birth control, and even
killing, of those considered "bad stock." Another example of an extension of evolutionary theory that's now widely regarded as unwarranted is "
Social Darwinism," a term given to the 19th century
Whig Malthusian theory developed by
Herbert Spencer into ideas about "
survival of the fittest" in commerce and human societies as a whole, and by others into claims that
social inequality, racism, and
imperialism were justified. However, contemporary scientists and philosophers consider these ideas to have been neither mandated by evolutionary theory nor supported by data.
Applications in technology
A major technological application of evolution is
artificial selection, which is the intentional selection of certain traits in a population of organisms. Humans have used artificial selection for thousands of years in the
domestication of plants and animals. More recently, such selection has become a vital part of
genetic engineering, with
selectable markers such as antibiotic resistance genes being used to manipulate DNA in
molecular biology.
As evolution can produce highly optimized processes and networks, it has many applications in
computer science. Here, simulations of evolution using
evolutionary algorithms and
artificial life started with the work of Nils Aall Barricelli in the 1960s, and was extended by
Alex Fraser, who published a series of papers on simulation of
artificial selection.
Artificial evolution became a widely recognized optimization method as a result of the work of
Ingo Rechenberg in the 1960s and early 1970s, who used
evolution strategies to solve complex engineering problems.
Genetic algorithms in particular became popular through the writing of
John Holland. As academic interest grew, dramatic increases in the power of computers allowed practical applications, including the 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 optimize the design of systems.
Further Information
Get more info on 'Theory Of Evolution'.
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