Introduction
Evolution is the process of change in all forms of life over generations, and evolutionary biology is the
study of how evolution occurs. Biological populations evolve through genetic changes that correspond
to changes in the organisms' observable traits. Genetic changes include mutations, which are caused by
damage or replication errors in organisms' DNA. As the genetic variation of a population drifts randomly
over generations, natural selection gradually leads traits to become more or less common based on the
relative reproductive success of organisms with those traits.
The "Paleontological Tree of the Vertebrates," from the 5th edition of The Evolution of Man (London,
1910) by Ernst Haeckel. The evolutionary history of species has been described as a tree, with many
branches arising from a single trunk.
The age of the Earth is about 4.5 billion years. The earliest undisputed evidence of life on Earth dates at
least from 3.5 billion years [Link] does not attempt to explain the origin of life (covered instead
by abiogenesis), but it does explain how early lifeforms evolved into the complex ecosystem that we see
today Based on the similarities between all present-day organisms, all life on Earth is assumed to have
originated through common descent from a last universal ancestor from which all known species have
diverged through the process of evolution.
All individuals have hereditary material in the form of genes received from their parents, which they
pass on to any offspring. Among offspring there are variations of genes due to the introduction of new
genes via random changes called mutations or via reshuffling of existing genes during sexual
[Link] offspring differs from the parent in minor random ways. If those differences are
helpful, the offspring is more likely to survive and reproduce. This means that more offspring in the next
generation will have that helpful difference and individuals will not have equal chances
of reproductive success. In this way, traits that result in organisms being better adapted to their living
conditions become more common in descendant [Link] differences accumulate resulting in
changes within the population. This process is responsible for the many diverse life forms in the world.
The modern understanding of evolution began with the 1859 publication of Charles Darwin's On the
Origin of Species. In addition, Gregor Mendel's work with plants helped to explain the hereditary
patterns of genetics. Fossil discoveries in palaeontology, advances in population genetics and a global
network of scientific research have provided further details into the mechanisms of evolution. Scientists
now have a good understanding of the origin of new species (speciation) and have observed the
speciation process in the laboratory and in the wild. Evolution is the principal scientific
theory that biologists use to understand life and is used in many disciplines,
including medicine, psychology, conservation biology, anthropology, forensics, agriculture and
other social-cultural applications.
Simple overview
The main ideas of evolution may be summarised as follows:
Life forms reproduce and therefore have a tendency to become more numerous.
Factors such as predation and competition work against the survival of individuals.
Each offspring differs from their parent(s) in minor, random ways.
If these differences are beneficial, the offspring is more likely to survive and reproduce.
This makes it likely that more offspring in the next generation will have beneficial differences and fewer
will have detrimental differences.
These differences accumulate over generations, resulting in changes within the population.
Over time, populations can split or branch off into new species.
These processes, collectively known as evolution, are responsible for the many diverse life forms seen in
the world.
Natural selection
Charles Darwin proposed the theory of evolution by natural selection.
Natural selection
In the 19th century, natural history collections and museums were popular. The European expansion
and naval expeditions employed naturalists, while curators of grand museums showcased preserved and
live specimens of the varieties of life. Charles Darwin was an English graduate educated and trained in
the disciplines of natural history. Such natural historians would collect, catalogue, describe and study
the vast collections of specimens stored and managed by curators at these museums. Darwin served as
a ship's naturalist on board HMS Beagle, assigned to a five-year research expedition around the world.
During his voyage, he observed and collected an abundance of organisms, being very interested in the
diverse forms of life along the coasts of South America and the neighbouring Galápagos Islands.
Darwin noted that orchids have complex adaptations to ensure pollination, all derived from basic floral
parts.
Darwin gained extensive experience as he collected and studied the natural history of life forms from
distant places. Through his studies, he formulated the idea that each species had developed from
ancestors with similar features. In 1838, he described how a process he called natural selection would
make this happen.
The size of a population depends on how much and how many resources are able to support it. For the
population to remain the same size year after year, there must be an equilibrium, or balance between
the population size and available resources. Since organisms produce more offspring than their
environment can support, not all individuals can survive out of each generation. There must be a
competitive struggle for resources that aid in survival. As a result, Darwin realised that it was not chance
alone that determined survival. Instead, survival of an organism depends on the differences of each
individual organism, or "traits," that aid or hinder survival and reproduction. Well-adapted individuals
are likely to leave more offspring than their less well-adapted competitors. Traits that hinder survival
and reproduction would disappear over generations. Traits that help an organism survive and reproduce
would accumulate over generations. Darwin realised that the unequal ability of individuals to survive
and reproduce could cause gradual changes in the population and used the term natural selection to
describe this process.
Observations of variations in animals and plants formed the basis of the theory of natural selection. For
example, Darwin observed that orchids and insects have a close relationship that allows
the pollination of the plants. He noted that orchids have a variety of structures that attract insects, so
that pollen from the flowers gets stuck to the insects' bodies. In this way, insects transport the pollen
from a male to a female orchid. In spite of the elaborate appearance of orchids, these specialised parts
are made from the same basic structures that make up other flowers. In his book, Fertilisation of
Orchids (1862), Darwin proposed that the orchid flowers were adapted from pre-existing parts, through
natural selection.
Darwin was still researching and experimenting with his ideas on natural selection when he received a
letter from Alfred Russel Wallace describing a theory very similar to his own. This led to an immediate
joint publication of both theories. Both Wallace and Darwin saw the history of life like a family tree, with
each fork in the tree's limbs being a common ancestor. The tips of the limbs represented modern
species and the branches represented the common ancestors that are shared amongst many different
species. To explain these relationships, Darwin said that all living things were related, and this meant
that all life must be descended from a few forms, or even from a single common ancestor. He called this
process descent with modification.
Darwin published his theory of evolution by natural selection in On the Origin of Species in [Link]
theory means that all life, including humanity, is a product of continuing natural processes. The
implication that all life on Earth has a common ancestor has met with objections from some religious
groups. Their objections are in contrast to the level of support for the theory by more than 99 percent of
those within the scientific community today.
Natural selection is commonly equated with survival of the fittest, but this expression originated
in Herbert Spencer's Principles of Biology in 1864, five years after Charles Darwin published his original
works. Survival of the fittest describes the process of natural selection incorrectly, because natural
selection is not only about survival and it is not always the fittest that survives.
Source of variation
Darwin's theory of natural selection laid the groundwork for modern evolutionary theory, and his
experiments and observations showed that the organisms in populations varied from each other, that
some of these variations were inherited, and that these differences could be acted on by natural
selection. However, he could not explain the source of these variations. Like many of his predecessors,
Darwin mistakenly thought that heritable traits were a product of use and disuse, and that features
acquired during an organism's lifetime could be passed on to its offspring. He looked for examples, such
as large ground feeding birds getting stronger legs through exercise, and weaker wings from not flying
until, like the ostrich, they could not fly at all.[21] This misunderstanding was called the inheritance of
acquired characters and was part of the theory of transmutation of species put forward in 1809 by Jean-
Baptiste Lamarck. In the late 19th century this theory became known as Lamarckism. Darwin produced
an unsuccessful theory he called pangenesis to try to explain how acquired characteristics could be
inherited. In the 1880s August Weismann's experiments indicated that changes from use and disuse
could not be inherited, and Lamarckism gradually fell from favour.
The missing information needed to help explain how new features could pass from a parent to its
offspring was provided by the pioneering genetics work of Gregor Mendel. Mendel's experiments with
several generations of pea plants demonstrated that inheritance works by separating and reshuffling
hereditary information during the formation of sex cells and recombining that information during
fertilisation. This is like mixing different hands of playing cards, with an organism getting a random mix
of half of the cards from one parent, and half of the cards from the other. Mendel called the
information factors; however, they later became known as genes. Genes are the basic units of heredity
in living organisms. They contain the information that directs the physical development and behaviour
of organisms.
Genes are made of DNA. DNA is a long molecule made up of individual molecules called nucleotides.
Genetic information is encoded in the sequence of nucleotides, that make up the DNA, just as the
sequence of the letters in words carries information on a page. The genes are like short instructions built
up of the "letters" of the DNA alphabet. Put together, the entire set of these genes gives enough
information to serve as an "instruction manual" of how to build and run an organism. The instructions
spelled out by this DNA alphabet can be changed, however, by mutations, and this may alter the
instructions carried within the genes. Within the cell, the genes are carried in chromosomes, which are
packages for carrying the DNA. It is the reshuffling of the chromosomes that results in unique
combinations of genes in offspring. Since genes interact with one another during the development of an
organism, novel combinations of genes produced by sexual reproduction can increase the genetic
variability of the population even without new mutations.[23] The genetic variability of a population can
also increase when members of that population interbreed with individuals from a different population
causing gene flow between the populations. This can introduce genes into a population that were not
present before.
Evolution is not a random process. Although mutations in DNA are random, natural selection is not a
process of chance: the environment determines the probability of reproductive success. Evolution is an
inevitable result of imperfectly copying, self-replicating organisms reproducing over billions of years
under the selective pressure of the environment. The outcome of evolution is not a perfectly designed
organism. The end products of natural selection are organisms that are adapted to their present
environments. Natural selection does not involve progress towards an ultimate goal. Evolution does not
strive for more advanced, more intelligent, or more sophisticated life forms.[25] For
example, fleas (wingless parasites) are descended from a winged, ancestral scorpionfly,
and snakes are lizards that no longer require limbs—although pythons still grow tiny structures that are
the remains of their ancestor's hind legs. Organisms are merely the outcome of variations that succeed
or fail, dependent upon the environmental conditions at the time.
Rapid environmental changes typically cause extinction. Of all species that have existed on Earth,
99.9 percent are now extinct. Since life began on Earth, five major mass extinctions have led to large and
sudden drops in the variety of species. The most recent, the Cretaceous–Paleogene extinction event,
occurred 66 million years ago.
Genetic drift
Genetic drift is a cause of allelic frequency change within populations of a species. Alleles are different
variations of specific genes. They determine things like hair colour, skin tone, eye colour and blood type;
in other words, all the genetic traits that vary between individuals. Genetic drift does not introduce new
alleles to a population, but it can reduce variation within a population by removing an allele from the
gene pool. Genetic drift is caused by random sampling of alleles. A truly random sample is a sample in
which no outside forces affect what is selected. It is like pulling marbles of the same size and weight but
of different colours from a brown paper bag. In any offspring, the alleles present are samples of the
previous generations alleles, and chance plays a role in whether an individual survives to reproduce and
to pass a sample of their generation onward to the next. The allelic frequency of a population is the ratio
of the copies of one specific allele that share the same form compared to the number of all forms of the
allele present in the population.
Genetic drift affects smaller populations more than it affects larger populations.[
Hardy–Weinberg principle
The Hardy–Weinberg principle states that under certain idealised conditions, including the absence of
selection pressures, a large population will have no change in the frequency of alleles as generations
pass.A population that satisfies these conditions is said to be in Hardy–Weinberg equilibrium. In
particular, Hardy and Weinberg showed that dominant and recessive alleles do not automatically tend
to become more and less frequent respectively, as had been thought previously.
The conditions for Hardy-Weinberg equilibrium include that there must be no mutations, immigration,
or emigration, all of which can directly change allelic frequencies. Additionally, mating must be totally
random, with all males (or females in some cases) being equally desirable mates. This ensures a true
random mixing of alleles. A population that is in Hardy–Weinberg equilibrium is analogous to a deck of
cards; no matter how many times the deck is shuffled, no new cards are added and no old ones are
taken away. Cards in the deck represent alleles in a population's gene pool.
In practice, no population can be in perfect Hardy-Weinberg equilibrium. The population's finite size,
combined with natural selection and many other effects, cause the allelic frequencies to change over
time.
Population bottleneck
Model of population bottleneck illustrates how alleles can be lost.
A population bottleneck occurs when the population of a species is reduced drastically over a short
period of time due to external forces.[35] In a true population bottleneck, the reduction does not favour
any combination of alleles; it is totally random chance which individuals survive. A bottleneck can reduce
or eliminate genetic variation from a population. Further drift events after the bottleneck event can also
reduce the population's genetic diversity. The lack of diversity created can make the population at risk to
other selective pressures.
A common example of a population bottleneck is the Northern elephant seal. Due to excessive hunting
throughout the 19th century, the population of the northern elephant seal was reduced to 30
individuals or less. They have made a full recovery, with the total number of individuals at around
100,000 and growing. The effects of the bottleneck are visible, however. The seals are more likely to
have serious problems with disease or genetic disorders, because there is almost no diversity in the
population.
Founder effect
In the founder effect, small new populations contain different allele frequencies from the parent
population.
The founder effect occurs when a small group from one population splits off and forms a new
population, often through geographic isolation. This new population's allelic frequency is probably
different from the original population's, and will change how common certain alleles are in the
populations. The founders of the population will determine the genetic makeup, and potentially the
survival, of the new population for generations
One example of the founder effect is found in the Amish migration to Pennsylvania in 1744. Two of the
founders of the colony in Pennsylvania carried the recessive allele for Ellis–van Creveld syndrome.
Because the Amish tend to be religious isolates, they interbreed, and through generations of this
practice the frequency of Ellis–van Creveld syndrome in the Amish people is much higher than the
frequency in the general population.[38]
Modern synthesis
The modern evolutionary synthesis is based on the concept that populations of organisms have
significant genetic variation caused by mutation and by the recombination of genes during sexual
reproduction. It defines evolution as the change in allelic frequencies within a population caused by
genetic drift, gene flow between sub populations, and natural selection. Natural selection is emphasised
as the most important mechanism of evolution; large changes are the result of the gradual accumulation
of small changes over long periods of time.
The modern evolutionary synthesis is the outcome of a merger of several different scientific fields to
produce a more cohesive understanding of evolutionary theory. In the 1920s, Ronald Fisher, J.B.S.
Haldane and Sewall Wright combined Darwin's theory of natural selection with statistical models
of Mendelian genetics, founding the discipline of population genetics. In the 1930s and 1940s, efforts
were made to merge population genetics, the observations of field naturalists on the distribution of
species and sub species, and analysis of the fossil record into a unified explanatory model. The
application of the principles of genetics to naturally occurring populations, by scientists such
as Theodosius Dobzhansky and Ernst Mayr, advanced the understanding of the processes.