Evolution

This article is about evolution in the biological sense. For other possible meanings, see Evolution (disambiguation).

Evolution is any process of growth or development that entails change. Most commonly, this refers to biological evolution, but evolution (disambiguation) for other uses. Evolution may be defined as the change in the frequency of an allele within a gene pool, caused by natural selection and/or genetic drift. Such changes over, long periods of time lead to major changes in phenotype. Important to evolution are other processes such as speciation.

Often it is shorthand for the modern theory of evolution of species based upon Darwin's idea of natural selection .

The current dominant theory of evolution is known as the "modern evolutionary synthesis" (or simply "modern synthesis"), referring to the synthesis of Darwin's theory of evolution by natural selection and Mendel's theory of the gene. According to this theory, the fundamental event of speciation is the genetic isolation of two populations, which allows their gene pools to diverge. Since the modern synthesis , biological evolution has been defined as changes in allele frequencies in a population from one generation to another. The remainder of this article addresses biological evolution.

Scientific theory

The commonly accepted scientific theory about how life has changed since it originated has three major aspects.

1. The common descent of all organisms from (more or less) a single ancestor.
2. The origin of novel traits in a lineage.
3. The mechanisms that cause some traits to persist while others perish.

Ancestry of organisms

Most biologists believe in common descent: that all life on Earth is descended from one common ancestor. This conclusion is based upon the fact that many traits of living organisms, such as the genetic code, seem arbitrary yet are shared by all organisms. Some have suggested that life may have had more than one origin, but the high degree of commonality argues strongly against multiple origins.

The study of the ancestry of species is phylogeny. Phylogeny has revealed that organs with radically different internal structures can bear a superficial resemblance and perform similar functions. These examples of analogous structures show that there are multiple ways to solve most problems and make it difficult to believe that the universal traits of life are all necessary. Likewise other organs with similar internal structures will perform radically different functions. Vertebrate limbs are a favorite example of homologous structures, organs on two organisms that share a basic structure that had existed in the last common ancestor of the organisms.

Further evidence of the universal ancestry of life is that abiogenesis has never been observed under controlled conditions, indicating that the origin of life from non-life, is either very rare or only happens under conditions that are not at all like those of modern earth.

Fossil evidence

Fossil evidence of prehistoric organisms has been found all over the Earth. The age of fossils can often be deduced, based upon the geologic context in which they are found. Some fossils show a resemblance to organisms alive today, while others are radically different. Fossils have been used to determine at what time a lineage developed, and can be used to demonstrate the continuity between two different lineages. Paleontologists investigate evolution largely through analysis of fossils.

The emergence of novel traits

If life is to change, then new traits must emerge at some point. Geneticists have studied how traits emerge and are passed to succeeding generations. In Darwin's time, there was no widely accepted in-depth mechanism for heritability. Today most inherited variation is traced to discrete, persistent entities called "genes". Genes are encoded in linear molecules called DNA. Changes in DNA are commonly called mutations. Furthermore, DNA variants may have little phenotypic effect in isolation but create new traits when combined in an organism through genetic recombination. Genetic recombination is produced both by the fusion of cells of opposite mating types (such as human sex), and by the transfer of material into an intact cell (such as bacterial conjugation and transformation).

Researchers are also investigating heritable variation that is not connected to variations in DNA sequence that influence standard DNA replication. The processes that produce this variation leave the genetic information intact and are often reversible. These are often referred to as epigenetic inheritance and may include phenomenon such as DNA methylation, prions, and structural inheritance. Investigations continue into whether these mechanisms allow for the production of specific beneficial heritable variation in response to environmental signals. If this is shown to be the case, then some instances of evolution would lie outside of the framework that Darwin established, which avoided any connection between environmental signals and the production of heritable variation. In general, Darwin knew little about the nature or source of heritable variation.

In addition to the mechanisms described above, the origin of novel traits may also be attributable to self-organizing properties at the level of the physics and chemistry of the organism (which some hold to be a violation of "strict" Darwinism). Self-organization in this context would refer to traits that were not directly encoded in the genome but rather would always expected to be present in a wide class of particular biological systems (see the section Neo-structuralist themes in evolutionary theory in the current article). In this view, most cogently expressed by Stuart Kauffman, natural selection "selects" only particular classes of systems, which happen to include systems which generate such "order for free" (Kauffman also calls this property "anti-chaos"). Several specific mechanisms to enable "order for free" such as the robustness of genetic regulatory networks, the spontaneous self-sustaining order of chemical reactions as autocatalytic sets and the properties of the RNA genotype-to-phenotype map (in this case, the RNA-sequence-to-RNA-shape mapping), have been cautiously incorporated as part of a workable theory as it applies to evolution. However, the entire program as outlined by Kauffman remains a matter for debate.

The foregoing potential sources of novel traits are not mutually exclusive, and most biologists would accept that each mechanism discussed has been demonstrated to be a possible way to generate such traits; however each would most likely assign different degrees of importance to each of the different mechanisms.

Microevolution and macroevolution

Main articles: Microevolution, Macroevolution

Microevolution refers to small-scale changes in gene-frequencies in a population over a few generations (population genetics is the branch of biology that provides the mathematical structure for the study of the process of microevolution). These changes may be due to a number of processes: mutation, gene flow, genetic drift, as well as natural selection. Macroevolution refers to large-scale changes in gene-frequencies in a population over a long period of time (and may culminate in the evolution of new species). The difference between the two is hard to distinguish because, over time, successive tiny mutations like those evidenced in microevolution could build up in isolated populations and eventually create entirely new genera, which is known as macroevolution. While microevolution has been demonstrated in the laboratory to the satisfaction of most observers, macroevolution has to be inferred from the fossil record and the traits of extant organisms. Its precise mechanisms are an active topic of discussion amongst scientists.

Differential survival of traits

Differential survival of characteristics that arise in the population means that some will become more frequent while others may be lost. Two processes are generally thought to contribute to the survival of a characteristic:

• Natural selection
• Genetic drift

Natural selection

Main article: Natural selection

Darwinism and its descendant theories state that biological evolution results through natural selection. Since natural selection is so important to Darwinism and modern theories of evolution, a very short summary of its main points follows:

• Organisms have children that inherit genes from their parents. These genes code for different characteristics in a person. Genetically, a child has 50% the DNA of each parent. However, depending on how the genotypes are inherited, the phenotypes may be manifested in different ways. The genotype is the basic code of the gene, and the phenotype is what is expressed in the individual. Two brown-eyed parents may be heterozygous for the eye color allele and end up having a child with the blue eyed phenotype. In plain English, kids are like mom and dad, though the mechanisms through which this occurs can get very complicated.
• Organisms have differing reproductive (sexual) success based on their traits in a given environment. In plain English, animals (or plants) that are good at what they do are more likely to survive and have kids.
• Therefore, over time, the types of organisms that have traits better adapted to their environment will tend to become the dominant ones in an environment, while organisms poorly adapted to their environment will become extinct.

Natural selection also provides for a mechanism by which life can sustain itself over time. Since, in the long run, environments always change, if successive generations did not develop adaptations which allowed them to survive and reproduce, species would simply die out as their biological niches die out. Therefore, life is allowed to persist over great spans of time, in the form of evolving species. The central role of natural selection in evolutionary theory has created a strong connection between that field and the study of ecology. The probable mutation effect is the proposition that a gene that is not under selection will be destroyed by accumulated mutations. This is an aspect of genome degradation.

Genetic drift

Main article: Genetic drift

Genetic drift describes changes in gene frequency that cannot be ascribed to selective pressures, but are due instead to events that are unrelated to inherited traits. This is especially important in small mating populations, which simply cannot have enough offspring to maintain the same gene distribution as the parental generation. Such fluctuations in gene frequency between successive generations may result in some genes disappearing from the population. Two separate populations that begin with the same gene frequency might, therefore, "drift" by random fluctuation into two divergent populations with different gene sets (i.e. genes that are present in one have been lost in the other). Rare sporadic events (volcanic explosion, meteor impact, etc.) might contribute to genetic drift by altering the gene frequency outside of "normal" selective pressures.

Recent developments in evolutionary theory

Symbiogenesis

Main article: Symbiogenesis

Another extension to the standard modern synthesis, advocated by Lynn Margulis, is symbiogenesis. Symbiogenesis argues that acquisition and accumulation of random mutations or genetic drift are not sufficient to explain how new inherited variations occur in evolution. This theory states that species arise from the merger of independent organisms through symbiosis. Symbiogenesis emphasizes the impact of co-operation rather than Darwinian competition. This commonly occurs in multigenomic organisms throughout nature.

Neo-structuralist themes in evolutionary theory

In the 1980s and 1990s there was a renewal of structuralist themes in evolutionary biology by biologists such as Brian Goodwin, that incorporates ideas from cybernetics and systems theory, and that emphasizes the role of self-organized processes as being at least as important as the role of natural selection. Some extreme variants consider natural selection as the result of biological evolution and not its cause, though most neo-structuralist biologists would not go this far.

The evolution of altruism

Main article: Altruism

Altruism has been one of the last (and most deeply embedded) thorns in the side of evolutionary theory, but recent developments in game theory have suggested explanations with an evolutionary context. If humans evolved, then so did human minds, and if minds evolved, then so does behaviour - including, according to these models, altruistic tendencies.

Theories of eusociality and the undoubted advantages of kin selection have made good progress in this direction, but they are far from unproblematic. Some writers have pointed out that the conscience is just another aspect of our mental behaviour, and propose an evolutionary explanation for the existence of conscience and therefore altruism. One recent suggestion, expressed most eloquently by the philosopher Daniel Dennett , was initially developed when considering the problem of so-called 'free riders' in the tragedy of the commons, a larger-scale version of the Prisoner's Dilemma.

An interesting example of altruism is found in the cellular slime molds, such as Dictyostelium mucoroides. These protists live as individual amoebae until starved, at which point they aggregate and form a multicellular fruiting body in which some cells sacrifice themselves to promote the survival of other cells in the fruiting body.

History of evolutionary thought

Present status

When talking about biological evolution, there is often confusion about the question of whether or not modern organisms have evolved (and are continuing to change) from older ancestral organisms and there are questions about the mechanism of the observed changes.

It is a fact that the frequency of traits changes in populations of organisms. Most biologists believe that this process can account for the differences between existing species, but the relative importance of the various mechanisms continues to be debated. The commonly accepted scientific theory today is known as modern synthesis (or the Neo-Darwinian synthesis), based primarily on Charles Darwin's theory of natural selection, but updated with newer discoveries in biology and genetics, in particular Mendelian inheritance. Population genetics is the branch of biology that provides the mathematical structure for the modern synthesis.

In popular usage, "the" theory of evolution refers to this or other Darwinian theories. However, within this framework there are still differences of opinion, for example between punctuated equilibrium and strict gradualism or regarding the relative importance of natural selection and genetic drift.

History evolutionary thought

Main article: History of evolutionary thought

The idea of biological evolution has existed since ancient times, but the modern theory wasn't established until the 18th and 19th centuries, with scientists such as Lamarck and Charles Darwin. Darwin greatly emphasized the difference between his two main inputs: establishing the fact of evolution, and proposing a theory, natural selection, to explain the mechanism of evolution.

As science has uncovered more and more information about the basic operations of life, such as genetics and molecular biology, theories of evolution have changed. The general trend has been not to overturn well-supported theories, but to supplant them with more detailed and therefore more complex ones.

While transmutation was accepted by a sizeable number of scientists before 1859, it was the publication of Charles Darwin's The Origin of Species which provided the first cogent mechanism by which evolutionary change could persist: his mechanism of natural selection. The evolutionary timeline outlines the major steps of evolution on Earth as expounded by this theory's proponents.

Following the dawn of molecular biology, it became clear that a major mechanism for variation within a population is the mutagenesis of DNA. An essential component to evolutionary theory is that during the cell cycle, DNA is copied very, but not entirely, faithfully. When these rare copying errors occur, they are said to introduce genetic mutations of three general consequences relative to the current environment: good, bad, or neutral. By definition, individuals with "good" mutations will be more likely to propagate, individuals with "bad" mutations will have less of a chance at successful reproduction, and those carrying "neutral" mutations will have neither an advantage nor a disadvantage. These definitions assume that the environment remains stable. Considered at the level of a single gene, these variations just described represent different genetic alleles. Following environmental change, alleles may retain their classification of good, bad, or neutral, or may shift into one of the other categories. Individuals carrying alleles formerly classified as neutral may now be "good" as they bear favourably adaptive mutations. Since neutral alleles can accumulate in the population without consequence while an environment is stable, they create a considerable reservoir for adaptability.

De Chardin's and Huxley's theories

Pierre Teilhard de Chardin and Julian Huxley formulated theories describing the gradual development of the Universe from subatomic particles to human society, considered by Teilhard as the last stage. (see Gaia theory). These are not generally recognized as scientifically rigorous.

Nine levels are described (scheme (http://noosphere.cc/evolutiontendencies.html)), the "classical" biological stages being levels 6, 7 & 8 of the universal evolution. Stages 1 to 5 are grouped into the Lithosphere, also called Geosphere or Physiosphere, where (the progress of) the structure of the organisms is ruled by structure, mechanical laws and coincidence. Stages 6 to 8 are grouped into the Biosphere, where (the progress of) the structure of the organisms is ruled by genetical mechanisms. The actual stage, stage 9, is called the Noosphere, where (the progress of) the structure of human society (socialization) is ruled by psychological, informational and communicative processes.

Social effect of evolutionary theory

Main article: Social effect of evolutionary theory

As the scientific explanation of life's diversity has developed, it has displaced the explanations held by a significant portion of humanity. As the theory of evolution includes an explanation of humanity's origins, it has had a profound impact onhuman societies. Some social conservatives have vigorously opposed acceptance of the scientific explanation due to perceived religious implications. The theory of evolution by natural selection has also been adopted as a foundation for various ethical systems, such as social Darwinism, although scientists emphasize that their work is intended purely as a description of nature. The notion that humans share ancestors with other animals has also impacted how some persons view the relationship between humans and other species.

The theory of evolution by natural selection has also been incorporated into other fields of knowledge, creating hybrids such as evolutionary psychology and sociobiology.

Creationism and evolution

Main article: Creationism

Ever since Darwin provided the first cogent mechanism for evolution, religious fundamentalists have claimed that the theory is false. They posit that a supreme and supernatural being (God) intentionally created the universe as a whole, and each species in particular. This view is commonly referred to as "creationism".

In response to world-wide scientific acceptance of the evolutionary process as a fact, more moderate views have emerged where God only provides a divine spark to make evolution happen (Evolutionary Creationism). Christians, Jews and Muslims who reject evolution point to the Bible and the Quran to support their views and have offered what they believe to be proof of the impossibility of macroevolution in particular. Others claim that life shows evidence of intelligent design. These arguments and their presentation as science are not accepted by the mainstream scientific community. In spite of this, creationists in the United States have succeeded in convincing some state governments to give "equal time" to their views in the classroom, and that the theory of evolution should be subject to critical evaluation much more intense than that given to any other scientific theory.

(See for example Disclaimer Adopted by Oklahoma Textbook Commission (http://www.his.atr.jp/~ray/textbook/disclaimer.html). As of 2004, a very similar notice is pasted into biology textbooks in Alabama as well. See also Cambrian Explosion for a discussion of issues raised by the notice.)

Links and references

Bibliography

• The Origin of Species by Charles Darwin
• Darwin's Dangerous Idea by Daniel Dennett
• The Selfish Gene and The Blind Watchmaker by Richard Dawkins
• Not in our Genes by Richard Lewontin, Steven Rose and Leon J. Kamin
• The Beak of the Finch by Jonathan Weiner
• Full House by Stephen Jay Gould
• How the Mind Works by Steven Pinker

See also

• Argument from evolution
• Atavism
• Blind Variation and Selective Retention
• Common descent
• Convergent evolution
• Endosymbiosis
• Evolution of Homo sapiens
• Evolution of sex
• Evolutionary tree
• Fitness landscape
• Gradualism
• Irreducible complexity
• Macromutation
• Neutral theory of molecular evolution
• Niche construction
• Quantum evolution
• Quasispecies model
• Sexual selection
• Teratogenesis
• Virus evolution

People

Famous evolution researchers and popularizers include:

• Charles Darwin
• Erasmus Darwin
• Richard Dawkins
• Daniel Dennett
• Jared Diamond
• Theodosius Dobzhansky
• Ronald Fisher
• Douglas Futuyma
• Stephen Jay Gould
• W. D. Hamilton
• Steve Jones
• Richard Lewontin
• Ernst Mayr
• Gregor Mendel
• Steven Pinker
• John Maynard Smith
• Alfred Russel Wallace
• Edward Osborne Wilson