In biology, 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 sometimes produce 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 age of the Earth is about 4.5 billion years. The earliest undisputed evidence of life on Earth dates from at least 3.5 billion years ago. Evolution 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 reproduction. The 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 populations. These 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, between 1856 and 1863, 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

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 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 1859. His 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 been 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., "Effects of the increased Use and Disuse of Parts, as controlled by Natural Selection" 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. 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.