Evolution is the change in the inherited characteristics of biological populations over successive generations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.
All life on Earth is descended from a last universal ancestor that lived approximately 3.8 billion years ago. Repeated speciation and the divergence of life can be inferred from 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 scientific argument for the theory of evolution by means of natural selection. Evolution by natural selection is a process that is inferred from three facts about populations:
1) more offspring are produced than can possibly survive,
2) traits vary among individuals, leading to different rates of survival and reproduction, and
3) trait differences are heritable.
Thus, when members of a population die they are replaced by the progeny of parents that were better adapted to survive and reproduce in the environment in which natural selection took place. This process creates and preserves 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 various aspects of evolution by forming and testing hypotheses, constructing scientific theories, using observational data, and 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.
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-Socratic 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 actualisations 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. The Roman poet and philosopher Titus Lucretius Carus proposed the possibility of evolutionary changes of organisms. 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. Leonardo da Vinci simply wrote, “Motion is the cause of all life”.
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 full-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 established 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 ofanimals 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 led 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 Weismann’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”.