Natural selection

Natural selection

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Encyclopedia
Natural selection is the nonrandom process by which biologic traits
Trait (biology)
A trait is a distinct variant of a phenotypic character of an organism that may be inherited, environmentally determined or be a combination of the two...

 become either more or less common in a population
Population
A population is all the organisms that both belong to the same group or species and live in the same geographical area. The area that is used to define a sexual population is such that inter-breeding is possible between any pair within the area and more probable than cross-breeding with individuals...

 as a function of differential reproduction of their bearers. It is a key mechanism of evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

.

The genetic variation
Genetic variation
Genetic variation, variation in alleles of genes, occurs both within and among populations. Genetic variation is important because it provides the “raw material” for natural selection. Genetic variation is brought about by mutation, a change in a chemical structure of a gene. Polyploidy is an...

 within a population of organisms may cause some individuals to survive and reproduce more successfully than others. Factors that affect reproductive success are also important, an issue that Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

 developed in his ideas on sexual selection
Sexual selection
Sexual selection, a concept introduced by Charles Darwin in his 1859 book On the Origin of Species, is a significant element of his theory of natural selection...

.

Natural selection acts on the phenotype
Phenotype
A phenotype is an organism's observable characteristics or traits: such as its morphology, development, biochemical or physiological properties, behavior, and products of behavior...

, or the observable characteristics of an organism, but the genetic
Genetics
Genetics , a discipline of biology, is the science of genes, heredity, and variation in living organisms....

 (heritable) basis of any phenotype that gives a reproductive advantage will become more common in a population (see allele frequency
Allele frequency
Allele frequency or Gene frequency is the proportion of all copies of a gene that is made up of a particular gene variant . In other words, it is the number of copies of a particular allele divided by the number of copies of all alleles at the genetic place in a population. It can be expressed for...

). Over time, this process can result in adaptation
Adaptation
An adaptation in biology is a trait with a current functional role in the life history of an organism that is maintained and evolved by means of natural selection. An adaptation refers to both the current state of being adapted and to the dynamic evolutionary process that leads to the adaptation....

s that specialize population
Population
A population is all the organisms that both belong to the same group or species and live in the same geographical area. The area that is used to define a sexual population is such that inter-breeding is possible between any pair within the area and more probable than cross-breeding with individuals...

s for particular ecological niche
Ecological niche
In ecology, a niche is a term describing the relational position of a species or population in its ecosystem to each other; e.g. a dolphin could potentially be in another ecological niche from one that travels in a different pod if the members of these pods utilize significantly different food...

s and may eventually result in the emergence of new species
Speciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...

. In other words, natural selection is an important process (though not the only process) by which evolution takes place within a population of organisms. As opposed to artificial selection
Artificial selection
Artificial selection describes intentional breeding for certain traits, or combination of traits. The term was utilized by Charles Darwin in contrast to natural selection, in which the differential reproduction of organisms with certain traits is attributed to improved survival or reproductive...

, in which humans favor specific traits, in natural selection the environment acts as a sieve through which only certain variations can pass.

Natural selection is one of the cornerstones of modern biology
Biology
Biology is a natural science concerned with the study of life and living organisms, including their structure, function, growth, origin, evolution, distribution, and taxonomy. Biology is a vast subject containing many subdivisions, topics, and disciplines...

. The term was introduced by Darwin in his influential 1859 book On the Origin of Species, in which natural selection was described as analogous to artificial selection
Artificial selection
Artificial selection describes intentional breeding for certain traits, or combination of traits. The term was utilized by Charles Darwin in contrast to natural selection, in which the differential reproduction of organisms with certain traits is attributed to improved survival or reproductive...

, a process by which animals and plants with traits considered desirable by human breeders are systematically favored for reproduction. The concept of natural selection was originally developed in the absence of a valid theory of heredity
Heredity
Heredity is the passing of traits to offspring . This is the process by which an offspring cell or organism acquires or becomes predisposed to the characteristics of its parent cell or organism. Through heredity, variations exhibited by individuals can accumulate and cause some species to evolve...

; at the time of Darwin's writing, nothing was known of modern genetics. The union of traditional Darwinian evolution
Darwinism
Darwinism is a set of movements and concepts related to ideas of transmutation of species or of evolution, including some ideas with no connection to the work of Charles Darwin....

 with subsequent discoveries in classical
Classical genetics
Classical genetics consists of the technique and methodologies of genetics that predate the advent of molecular biology. A key discovery of classical genetics in eukaryotes was genetic linkage...

 and molecular genetics
Molecular genetics
Molecular genetics is the field of biology and genetics that studies the structure and function of genes at a molecular level. The field studies how the genes are transferred from generation to generation. Molecular genetics employs the methods of genetics and molecular biology...

 is termed the modern evolutionary synthesis
Modern evolutionary synthesis
The modern evolutionary synthesis is a union of ideas from several biological specialties which provides a widely accepted account of evolution...

. Natural selection remains the primary explanation for adaptive evolution.

General principles


Natural variation occurs among the individuals of any population of organisms. Many of these differences do not affect survival (such as differences in eye color in humans), but some differences may improve the chances of survival of a particular individual. A rabbit that runs faster than others may be more likely to escape from predators, and algae
Algae
Algae are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms, such as the giant kelps that grow to 65 meters in length. They are photosynthetic like plants, and "simple" because their tissues are not organized into the many...

 that are more efficient at extracting energy from sunlight will grow faster. Something that increases an animal's survival will often also include its reproductive rate; however, sometimes there is a trade-off between survival and current reproduction. Ultimately, what matters is total lifetime reproduction of the animal.

For example, the peppered moth
Peppered moth
The peppered moth is a temperate species of night-flying moth. Peppered moth evolution is often used by educators as an example of natural selection.- Distribution :...

 exists in both light and dark colors in the United Kingdom
United Kingdom
The United Kingdom of Great Britain and Northern IrelandIn the United Kingdom and Dependencies, other languages have been officially recognised as legitimate autochthonous languages under the European Charter for Regional or Minority Languages...

, but during the industrial revolution
Industrial Revolution
The Industrial Revolution was a period from the 18th to the 19th century where major changes in agriculture, manufacturing, mining, transportation, and technology had a profound effect on the social, economic and cultural conditions of the times...

 many of the trees on which the moths rested became blackened by soot, giving the dark-colored moths an advantage in hiding from predators. This gave dark-colored moths a better chance of surviving to produce dark-colored offspring, and in just fifty years from the first dark moth being caught, nearly all of the moths in industrial Manchester
Manchester
Manchester is a city and metropolitan borough in Greater Manchester, England. According to the Office for National Statistics, the 2010 mid-year population estimate for Manchester was 498,800. Manchester lies within one of the UK's largest metropolitan areas, the metropolitan county of Greater...

 were dark. The balance was reversed by the effect of the Clean Air Act 1956
Clean Air Act 1956
The Clean Air Act 1956 was an Act of the Parliament of the United Kingdom passed in response to London's Great Smog of 1952. It was in effect until 1964, and sponsored by the Ministry of Housing and Local Government in England and the Department of Health for Scotland.The Act introduced a number of...

, and the dark moths became rare again, demonstrating the influence of natural selection on peppered moth evolution
Peppered moth evolution
The evolution of the peppered moth over the last two hundred years has been studied in detail. Originally, the vast majority of peppered moths had light colouration, which effectively camouflaged them against the light-coloured trees and lichens which they rested upon...

.

If the traits that give these individuals a reproductive advantage are also heritable, that is, passed from parent to child, then there will be a slightly higher proportion of fast rabbits or efficient algae in the next generation. This is known as differential reproduction. Even if the reproductive advantage is very slight, over many generations any heritable advantage will become dominant in the population. In this way the natural environment of an organism "selects" for traits that confer a reproductive advantage, causing gradual changes or evolution of life. This effect was first described and named by Charles Darwin.

The concept of natural selection predates the understanding of genetics, the mechanism of heredity for all known life forms. In modern terms, selection acts on an organism's phenotype, or observable characteristics, but it is the organism's genetic make-up or genotype
Genotype
The genotype is the genetic makeup of a cell, an organism, or an individual usually with reference to a specific character under consideration...

 that is inherited. The phenotype is the result of the genotype and the environment in which the organism lives (see Genotype-phenotype distinction
Genotype-phenotype distinction
The genotype–phenotype distinction is drawn in genetics. "Genotype" is an organism's full hereditary information, even if not expressed. "Phenotype" is an organism's actual observed properties, such as morphology, development, or behavior...

).

This is the link between natural selection and genetics, as described in the modern evolutionary synthesis
Modern evolutionary synthesis
The modern evolutionary synthesis is a union of ideas from several biological specialties which provides a widely accepted account of evolution...

. Although a complete theory of evolution also requires an account of how genetic variation arises in the first place (such as by mutation
Mutation
In molecular biology and genetics, mutations are changes in a genomic sequence: the DNA sequence of a cell's genome or the DNA or RNA sequence of a virus. They can be defined as sudden and spontaneous changes in the cell. Mutations are caused by radiation, viruses, transposons and mutagenic...

 and sexual reproduction
Sexual reproduction
Sexual reproduction is the creation of a new organism by combining the genetic material of two organisms. There are two main processes during sexual reproduction; they are: meiosis, involving the halving of the number of chromosomes; and fertilization, involving the fusion of two gametes and the...

) and includes other evolutionary mechanisms (such as genetic drift and gene flow
Gene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...

), natural selection appears to be the most important mechanism for creating complex adaptations in nature.

Nomenclature and usage


The term natural selection has slightly different definitions in different contexts. It is most often defined to operate on heritable traits, because these are the traits that directly participate in evolution. However, natural selection is "blind" in the sense that changes in phenotype (physical and behavioral characteristics) can give a reproductive advantage regardless of whether or not the trait is heritable (non heritable traits can be the result of environmental factors or the life experience of the organism).

Following Darwin's primary usage the term is often used to refer to both the evolutionary consequence of blind selection and to its mechanisms. It is sometimes helpful to explicitly distinguish between selection's mechanisms and its effects; when this distinction is important, scientists define "natural selection" specifically as "those mechanisms that contribute to the selection of individuals that reproduce", without regard to whether the basis of the selection is heritable. This is sometimes referred to as "phenotypic natural selection".

Traits that cause greater reproductive success of an organism are said to be selected for, whereas those that reduce success are selected against. Selection for a trait may also result in the selection of other correlated traits that do not themselves directly influence reproductive advantage. This may occur as a result of pleiotropy
Pleiotropy
Pleiotropy occurs when one gene influences multiple phenotypic traits. Consequently, a mutation in a pleiotropic gene may have an effect on some or all traits simultaneously...

 or gene linkage.

Fitness



The concept of fitness
Fitness (biology)
Fitness is a central idea in evolutionary theory. It can be defined either with respect to a genotype or to a phenotype in a given environment...

 is central to natural selection. In broad terms, individuals that are more "fit" have better potential for survival, as in the well-known phrase "survival of the fittest
Survival of the fittest
"Survival of the fittest" is a phrase originating in evolutionary theory, as an alternative description of Natural selection. The phrase is today commonly used in contexts that are incompatible with the original meaning as intended by its first two proponents: British polymath philosopher Herbert...

". However, as with natural selection above, the precise meaning of the term is much more subtle, and Richard Dawkins
Richard Dawkins
Clinton Richard Dawkins, FRS, FRSL , known as Richard Dawkins, is a British ethologist, evolutionary biologist and author...

 manages in his later books to avoid it entirely. (He devotes a chapter of his book, The Extended Phenotype
The Extended Phenotype
The Extended Phenotype is a biological concept introduced by Richard Dawkins in a 1982 book with the same title. The main idea is that phenotype should not be limited to biological processes such as protein biosynthesis or tissue growth, but extended to include all effects that a gene has on its...

, to discussing the various senses in which the term is used). Modern evolutionary theory defines fitness not by how long an organism lives, but by how successful it is at reproducing. If an organism lives half as long as others of its species, but has twice as many offspring surviving to adulthood, its genes will become more common in the adult population of the next generation.

Though natural selection acts on individuals, the effects of chance mean that fitness can only really be defined "on average" for the individuals within a population. The fitness of a particular genotype corresponds to the average effect on all individuals with that genotype. Very low-fitness genotypes cause their bearers to have few or no offspring on average; examples include many human genetic disorder
Genetic disorder
A genetic disorder is an illness caused by abnormalities in genes or chromosomes, especially a condition that is present from before birth. Most genetic disorders are quite rare and affect one person in every several thousands or millions....

s like cystic fibrosis
Cystic fibrosis
Cystic fibrosis is a recessive genetic disease affecting most critically the lungs, and also the pancreas, liver, and intestine...

.

Since fitness is an averaged quantity, it is also possible that a favorable mutation arises in an individual that does not survive to adulthood for unrelated reasons. Fitness also depends crucially upon the environment. Conditions like sickle-cell anemia may have low fitness in the general human population, but because the sickle-cell trait confers immunity from malaria, it has high fitness value in populations that have high malaria infection rates.

Types of selection


Natural selection can act on any heritable phenotypic trait, and selective pressure can be produced by any aspect of the environment, including sexual selection
Sexual selection
Sexual selection, a concept introduced by Charles Darwin in his 1859 book On the Origin of Species, is a significant element of his theory of natural selection...

 and competition with members of the same or other species. However, this does not imply that natural selection is always directional and results in adaptive evolution; natural selection often results in the maintenance of the status quo by eliminating less fit variants.

The unit of selection
Unit of selection
A unit of selection is a biological entity within the hierarchy of biological organisation that is subject to natural selection...

 can be the individual or it can be another level within the hierarchy of biological organisation, such as genes, cells
Cell (biology)
The cell is the basic structural and functional unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often called the building block of life. The Alberts text discusses how the "cellular building blocks" move to shape developing embryos....

, and kin groups
Kin selection
Kin selection refers to apparent strategies in evolution that favor the reproductive success of an organism's relatives, even at a cost to the organism's own survival and reproduction. Charles Darwin was the first to discuss the concept of group/kin selection...

. There is still debate about whether natural selection acts at the level of groups or species
Group selection
In evolutionary biology, group selection refers to the idea that alleles can become fixed or spread in a population because of the benefits they bestow on groups, regardless of the alleles' effect on the fitness of individuals within that group....

 to produce adaptations that benefit a larger, non-kin group. Likewise, there is debate as to whether selection at the molecular level prior to gene mutations and fertilization of the zygote should be ascribed to conventional natural selection because traditionally natural selection is an environmental and exterior force that acts upon a phenotype typically after birth. Some science journalists distinguish gene selection from natural selection by informally referencing selection of mutations as "pre-selection."

Selection at a different level such as the gene can result in an increase in fitness for that gene, while at the same time reducing the fitness of the individuals carrying that gene, in a process called intragenomic conflict
Intragenomic conflict
The selfish gene theory postulates that natural selection will increase the frequency of those genes whose phenotypic effects ensure their successful replication...

. Overall, the combined effect of all selection pressures at various levels determines the overall fitness of an individual, and hence the outcome of natural selection.


Natural selection occurs at every life stage of an individual. An individual organism must survive until adulthood before it can reproduce, and selection of those that reach this stage is called viability selection. In many species, adults must compete with each other for mates via sexual selection, and success in this competition determines who will parent the next generation. When individuals can reproduce more than once, a longer survival in the reproductive phase increases the number of offspring, called survival selection.

The fecundity
Fecundity
Fecundity, derived from the word fecund, generally refers to the ability to reproduce. In demography, fecundity is the potential reproductive capacity of an individual or population. In biology, the definition is more equivalent to fertility, or the actual reproductive rate of an organism or...

 of both females and males (for example, giant sperm
Spermatozoon
A spermatozoon is a motile sperm cell, or moving form of the haploid cell that is the male gamete. A spermatozoon joins an ovum to form a zygote...

 in certain species of Drosophila
Drosophila
Drosophila is a genus of small flies, belonging to the family Drosophilidae, whose members are often called "fruit flies" or more appropriately pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species to linger around overripe or rotting fruit...

) can be limited via "fecundity selection". The viability of produced gamete
Gamete
A gamete is a cell that fuses with another cell during fertilization in organisms that reproduce sexually...

s can differ, while intragenomic conflicts such as meiotic drive between the haploid gametes can result in gametic or "genic selection". Finally, the union of some combinations of eggs and sperm might be more compatible than others; this is termed compatibility selection.

Sexual selection



It is useful to distinguish between "ecological selection
Ecological selection
Ecological selection refers to natural selection minus sexual selection, i.e. strictly ecological processes that operate on a species' inherited traits without reference to mating or secondary sex characteristics...

" and "sexual selection". Ecological selection covers any mechanism of selection as a result of the environment (including relatives, e.g. kin selection
Kin selection
Kin selection refers to apparent strategies in evolution that favor the reproductive success of an organism's relatives, even at a cost to the organism's own survival and reproduction. Charles Darwin was the first to discuss the concept of group/kin selection...

, competition
Competition (biology)
Competition is an interaction between organisms or species, in which the fitness of one is lowered by the presence of another. Limited supply of at least one resource used by both is required. Competition both within and between species is an important topic in ecology, especially community ecology...

, and infanticide
Infanticide (zoology)
In animals, infanticide involves the killing of young offspring by a mature animal of its own species, and is studied in zoology, specifically in the field of ethology. Ovicide is the analogous destruction of eggs. Although human infanticide has been widely studied, the practice has been observed...

), while "sexual selection" refers specifically to competition for mates.

Sexual selection can be intrasexual, as in cases of competition among individuals of the same sex in a population, or intersexual, as in cases where one sex controls reproductive access by choosing among a population of available mates. Most commonly, intrasexual selection involves male–male competition and intersexual selection involves female choice of suitable males, due to the generally greater investment of resources for a female than a male in a single offspring. However, some species exhibit sex-role reversed behavior in which it is males that are most selective in mate choice; the best-known examples of this pattern occur in some fishes of the family Syngnathidae
Syngnathidae
Syngnathidae is a family of fish which includes the seahorses, the pipefishes, and the weedy and leafy sea dragons. The name is derived from Greek, meaning "fused jaw" - syn meaning fused or together, and gnathus meaning jaws. This fused jaw trait is something the entire family has in common...

, though likely examples have also been found in amphibian and bird species.

Some features that are confined to one sex only of a particular species can be explained by selection exercised by the other sex in the choice of a mate, for example, the extravagant plumage of some male birds. Similarly, aggression between members of the same sex is sometimes associated with very distinctive features, such as the antlers of stag
STAG
STAG: A Test of Love is a reality TV show hosted by Tommy Habeeb. Each episode profiles an engaged couple a week or two before their wedding. The cameras then follow the groom on his bachelor party...

s, which are used in combat with other stags. More generally, intrasexual selection is often associated with sexual dimorphism
Sexual dimorphism
Sexual dimorphism is a phenotypic difference between males and females of the same species. Examples of such differences include differences in morphology, ornamentation, and behavior.-Examples:-Ornamentation / coloration:...

, including differences in body size between males and females of a species.

Examples of natural selection



A well-known example of natural selection in action is the development of antibiotic resistance
Antibiotic resistance
Antibiotic resistance is a type of drug resistance where a microorganism is able to survive exposure to an antibiotic. While a spontaneous or induced genetic mutation in bacteria may confer resistance to antimicrobial drugs, genes that confer resistance can be transferred between bacteria in a...

 in microorganism
Microorganism
A microorganism or microbe is a microscopic organism that comprises either a single cell , cell clusters, or no cell at all...

s. Since the discovery of penicillin
Penicillin
Penicillin is a group of antibiotics derived from Penicillium fungi. They include penicillin G, procaine penicillin, benzathine penicillin, and penicillin V....

 in 1928, antibiotic
Antibiotic
An antibacterial is a compound or substance that kills or slows down the growth of bacteria.The term is often used synonymously with the term antibiotic; today, however, with increased knowledge of the causative agents of various infectious diseases, antibiotic has come to denote a broader range of...

s have been used to fight bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...

l diseases. Natural populations of bacteria contain, among their vast numbers of individual members, considerable variation in their genetic material, primarily as the result of mutations. When exposed to antibiotics, most bacteria die quickly, but some may have mutations that make them slightly less susceptible. If the exposure to antibiotics is short, these individuals will survive the treatment. This selective elimination of maladapted individuals from a population is natural selection.

These surviving bacteria will then reproduce again, producing the next generation. Due to the elimination of the maladapted individuals in the past generation, this population contains more bacteria that have some resistance against the antibiotic. At the same time, new mutations occur, contributing new genetic variation to the existing genetic variation. Spontaneous mutations are very rare, and advantageous mutations are even rarer. However, populations of bacteria are large enough that a few individuals will have beneficial mutations. If a new mutation reduces their susceptibility to an antibiotic, these individuals are more likely to survive when next confronted with that antibiotic.

Given enough time and repeated exposure to the antibiotic, a population of antibiotic-resistant bacteria will emerge. This new changed population of antibiotic-resistant bacteria is optimally adapted to the context it evolved in. At the same time, it is not necessarily optimally adapted any more to the old antibiotic free environment. The end result of natural selection is two populations that are both optimally adapted to their specific environment, while both perform substandard in the other environment.

The widespread use and misuse of antibiotics has resulted in increased microbial resistance to antibiotics in clinical use, to the point that the methicillin-resistant Staphylococcus aureus
Methicillin-resistant Staphylococcus aureus
Methicillin-resistant Staphylococcus aureus is a bacterium responsible for several difficult-to-treat infections in humans. It is also called multidrug-resistant Staphylococcus aureus and oxacillin-resistant Staphylococcus aureus...

 (MRSA) has been described as a "superbug" because of the threat it poses to health and its relative invulnerability to existing drugs. Response strategies typically include the use of different, stronger antibiotics; however, new strains
Strain (biology)
In biology, a strain is a low-level taxonomic rank used in three related ways.-Microbiology and virology:A strain is a genetic variant or subtype of a micro-organism . For example, a "flu strain" is a certain biological form of the influenza or "flu" virus...

 of MRSA have recently emerged that are resistant even to these drugs.

This is an example of what is known as an evolutionary arms race
Evolutionary arms race
In evolutionary biology, an evolutionary arms race is an evolutionary struggle between competing sets of co-evolving genes that develop adaptations and counter-adaptations against each other, resembling an arms race, which are also examples of positive feedback...

, in which bacteria continue to develop strains that are less susceptible to antibiotics, while medical researchers continue to develop new antibiotics that can kill them. A similar situation occurs with pesticide resistance
Pesticide resistance
Pesticide resistance is the adaptation of pest population targeted by a pesticide resulting in decreased susceptibility to that chemical. In other words, pests develop a resistance to a chemical through natural selection: the most resistant organisms are the ones to survive and pass on their...

 in plants and insects. Arms races are not necessarily induced by man; a well-documented example involves the spread of a gene in the butterfly Hypolimnas bolina
Hypolimnas bolina
The Great Eggfly , also called the Blue Moon Butterfly in New Zealand or Common Eggfly, is a species of nymphalid butterfly.-Race bolina:...

 suppressing male-killing activity by Wolbachia
Wolbachia
Wolbachia is a genus of bacteria which infects arthropod species, including a high proportion of insects , as well as some nematodes. It is one of the world's most common parasitic microbes and is possibly the most common reproductive parasite in the biosphere...

 bacteria parasites on the island of Samoa, where the spread of the gene is known to have occurred over a period of just five years

Evolution by means of natural selection



A prerequisite for natural selection to result in adaptive evolution
Adaptation
An adaptation in biology is a trait with a current functional role in the life history of an organism that is maintained and evolved by means of natural selection. An adaptation refers to both the current state of being adapted and to the dynamic evolutionary process that leads to the adaptation....

, novel traits and speciation, is the presence of heritable genetic variation that results in fitness differences. Genetic variation is the result of mutations, recombination
Genetic recombination
Genetic recombination is a process by which a molecule of nucleic acid is broken and then joined to a different one. Recombination can occur between similar molecules of DNA, as in homologous recombination, or dissimilar molecules, as in non-homologous end joining. Recombination is a common method...

s and alterations in the karyotype
Karyotype
A karyotype is the number and appearance of chromosomes in the nucleus of an eukaryotic cell. The term is also used for the complete set of chromosomes in a species, or an individual organism.p28...

 (the number, shape, size and internal arrangement of the chromosome
Chromosome
A chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...

s). Any of these changes might have an effect that is highly advantageous or highly disadvantageous, but large effects are very rare. In the past, most changes in the genetic material were considered neutral or close to neutral because they occurred in noncoding DNA
Noncoding DNA
In genetics, noncoding DNA describes components of an organism's DNA sequences that do not encode for protein sequences. In many eukaryotes, a large percentage of an organism's total genome size is noncoding DNA, although the amount of noncoding DNA, and the proportion of coding versus noncoding...

 or resulted in a synonymous substitution
Synonymous substitution
A synonymous substitution is the evolutionary substitution of one base for another in an exon of a gene coding for a protein, such that the produced amino acid sequence is not modified. Synonymous substitutions and mutations affecting noncoding DNA are collectively known as silent mutations...

. However, recent research suggests that many mutations in non-coding DNA do have slight deleterious effects. Although both mutation rates and average fitness effects of mutations are dependent on the organism, estimates from data in human
Human
Humans are the only living species in the Homo genus...

s have found that a majority of mutations are slightly deleterious.

By the definition of fitness, individuals with greater fitness are more likely to contribute offspring to the next generation, while individuals with lesser fitness are more likely to die early or fail to reproduce. As a result, alleles that on average result in greater fitness become more abundant in the next generation, while alleles that in general reduce fitness become rarer. If the selection forces remain the same for many generations, beneficial alleles become more and more abundant, until they dominate the population, while alleles with a lesser fitness disappear. In every generation, new mutations and re-combinations arise spontaneously, producing a new spectrum of phenotypes. Therefore, each new generation will be enriched by the increasing abundance of alleles that contribute to those traits that were favored by selection, enhancing these traits over successive generations.

Some mutations occur in so-called regulatory genes
Regulatory sequence
A regulatory sequence is a segment of DNA where regulatory proteins such as transcription factors bind preferentially. These regulatory proteins bind to short stretches of DNA called regulatory regions, which are appropriately positioned in the genome, usually a short distance 'upstream' of the...

. Changes in these can have large effects on the phenotype of the individual because they regulate the function of many other genes. Most, but not all, mutations in regulatory genes result in non-viable zygote
Zygote
A zygote , or zygocyte, is the initial cell formed when two gamete cells are joined by means of sexual reproduction. In multicellular organisms, it is the earliest developmental stage of the embryo...

s. Examples of nonlethal regulatory mutations occur in HOX genes
Homeobox
A homeobox is a DNA sequence found within genes that are involved in the regulation of patterns of anatomical development in animals, fungi and plants.- Discovery :...

 in humans, which can result in a cervical rib
Cervical rib
A cervical rib is a supernumerary rib which arises from the seventh cervical vertebra. It is a congenital abnormality located above the normal first rib. A cervical rib is present in only about 1 in 500 of people; in even rarer cases, an individual may have two cervical ribs...

 or polydactyly
Polydactyly
Polydactyly or polydactylism , also known as hyperdactyly, is a congenital physical anomaly in humans, dogs, and cats having supernumerary fingers or toes....

, an increase in the number of fingers or toes. When such mutations result in a higher fitness, natural selection will favor these phenotypes and the novel trait will spread in the population.
Established traits are not immutable; traits that have high fitness in one environmental context may be much less fit if environmental conditions change. In the absence of natural selection to preserve such a trait, it will become more variable and deteriorate over time, possibly resulting in a vestigial
Vestigial structure
Vestigiality describes homologous characters of organisms that have seemingly lost all or most of their original function in a species through evolution. These may take various forms such as anatomical structures, behaviors and biochemical pathways. Some of these disappear early in embryonic...

 manifestation of the trait, also called evolutionary baggage
Evolutionary baggage
Evolutionary baggage is the part of the genome of a population that was advantageous in past individuals but is disadvantageous under the pressures exerted by natural selection today.-Origin:...

. In many circumstances, the apparently vestigial structure may retain a limited functionality, or may be co-opted for other advantageous traits in a phenomenon known as preadaptation
Preadaptation
In evolutionary biology, preadaptation describes a situation where a species evolves to use a preexisting structure or trait inherited from an ancestor for a potentially unrelated function...

. A famous example of a vestigial structure, the eye
Eye
Eyes are organs that detect light and convert it into electro-chemical impulses in neurons. The simplest photoreceptors in conscious vision connect light to movement...

 of the blind mole rat
Blind mole rat
The genus Spalax contains the blind, fossorial, or subterranean mole rats, which are one of several types of rodents that are called mole rats. The hystricognath mole rats of the family Bathyergidae are completely unrelated, but some other forms are also in the family Spalacidae...

, is believed to retain function in photoperiod perception.

Speciation



Speciation
Speciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...

 requires selective mating, which result in a reduced gene flow
Gene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...

. Selective mating can be the result of 1. Geographic isolation, 2. Behavioral isolation, or 3. Temporal isolation. For example, a change in the physical environment (geographic isolation by an extrinsic barrier) would follow number 1, a change in camouflage for number 2 or a shift in mating times (i.e., one species of deer shifts location and therefore changes its "rut") for number 3.

Over time, these subgroups might diverge radically to become different species, either because of differences in selection pressures on the different subgroups, or because different mutations arise spontaneously in the different populations, or because of founder effect
Founder effect
In population genetics, the founder effect is the loss of genetic variation that occurs when a new population is established by a very small number of individuals from a larger population. It was first fully outlined by Ernst Mayr in 1942, using existing theoretical work by those such as Sewall...

s – some potentially beneficial alleles may, by chance, be present in only one or other of two subgroups when they first become separated. A lesser-known mechanism of speciation occurs via hybridization, well-documented in plants and occasionally observed in species-rich groups of animals such as cichlid
Cichlid
Cichlids are fishes from the family Cichlidae in the order Perciformes. Cichlids are members of a group known as the Labroidei along with the wrasses , damselfish , and surfperches . This family is both large and diverse. At least 1,300 species have been scientifically described, making it one of...

 fishes. Such mechanisms of rapid speciation can reflect a mechanism of evolutionary change known as punctuated equilibrium
Punctuated equilibrium
Punctuated equilibrium is a theory in evolutionary biology which proposes that most species will exhibit little net evolutionary change for most of their geological history, remaining in an extended state called stasis...

, which suggests that evolutionary change and in particular speciation typically happens quickly after interrupting long periods of stasis.

Genetic changes within groups result in increasing incompatibility between the genomes of the two subgroups, thus reducing gene flow between the groups. Gene flow will effectively cease when the distinctive mutations characterizing each subgroup become fixed. As few as two mutations can result in speciation: if each mutation has a neutral or positive effect on fitness when they occur separately, but a negative effect when they occur together, then fixation of these genes in the respective subgroups will lead to two reproductively isolated populations. According to the biological species concept, these will be two different species.

Historical development




Pre-Darwinian theories


Several ancient philosophers expressed the idea that nature produces a huge variety of creatures, randomly, and that only those creatures that manage to provide for themselves and reproduce successfully survive; well-known examples include Empedocles
Empedocles
Empedocles was a Greek pre-Socratic philosopher and a citizen of Agrigentum, a Greek city in Sicily. Empedocles' philosophy is best known for being the originator of the cosmogenic theory of the four Classical elements...

 and his intellectual successor, Lucretius
Lucretius
Titus Lucretius Carus was a Roman poet and philosopher. His only known work is an epic philosophical poem laying out the beliefs of Epicureanism, De rerum natura, translated into English as On the Nature of Things or "On the Nature of the Universe".Virtually no details have come down concerning...

, while related ideas were later refined by Aristotle
Aristotle
Aristotle was a Greek philosopher and polymath, a student of Plato and teacher of Alexander the Great. His writings cover many subjects, including physics, metaphysics, poetry, theater, music, logic, rhetoric, linguistics, politics, government, ethics, biology, and zoology...

. The struggle for existence was later described by Al-Jahiz
Al-Jahiz
Al-Jāḥiẓ was an Arabic prose writer and author of works of literature, Mu'tazili theology, and politico-religious polemics.In biology, Al-Jahiz introduced the concept of food chains and also proposed a scheme of animal evolution that entailed...

, who argued that environmental factors influence animals to develop new characteristics to ensure survival.

Abu Rayhan Biruni described the idea of artificial selection
Artificial selection
Artificial selection describes intentional breeding for certain traits, or combination of traits. The term was utilized by Charles Darwin in contrast to natural selection, in which the differential reproduction of organisms with certain traits is attributed to improved survival or reproductive...

 and argued that nature works in much the same way. Such classical arguments were reintroduced in the 18th century by Pierre Louis Maupertuis
Pierre Louis Maupertuis
Pierre-Louis Moreau de Maupertuis was a French mathematician, philosopher and man of letters. He became the Director of the Académie des Sciences, and the first President of the Berlin Academy of Science, at the invitation of Frederick the Great....

 and others, including Charles Darwin's grandfather Erasmus Darwin
Erasmus Darwin
Erasmus Darwin was an English physician who turned down George III's invitation to be a physician to the King. One of the key thinkers of the Midlands Enlightenment, he was also a natural philosopher, physiologist, slave trade abolitionist,inventor and poet...

. While these forerunners had an influence on Darwinism, they later had little influence on the trajectory of evolutionary thought after Charles Darwin.

Until the early 19th century, the prevailing view in Western societies was that differences between individuals of a species were uninteresting departures from their Platonic idealism (or typus) of created kinds. However, the theory of uniformitarianism
Uniformitarianism (science)
In the philosophy of naturalism, the uniformitarianism assumption is that the same natural laws and processes that operate in the universe now, have always operated in the universe in the past and apply everywhere in the universe. It has included the gradualistic concept that "the present is the...

 in geology
Geology
Geology is the science comprising the study of solid Earth, the rocks of which it is composed, and the processes by which it evolves. Geology gives insight into the history of the Earth, as it provides the primary evidence for plate tectonics, the evolutionary history of life, and past climates...

 promoted the idea that simple, weak forces could act continuously over long periods of time to produce radical changes in the Earth's landscape. The success of this theory raised awareness of the vast scale of geological time and made plausible the idea that tiny, virtually imperceptible changes in successive generations could produce consequences on the scale of differences between species.

Early 19th-century evolutionists such as Jean Baptiste Lamarck suggested the inheritance of acquired characteristics as a mechanism for evolutionary change; adaptive traits acquired by an organism during its lifetime could be inherited by that organism's progeny, eventually causing transmutation of species
Transmutation of species
Transmutation of species was a term used by Jean Baptiste Lamarck in 1809 for his theory that described the altering of one species into another, and the term is often used to describe 19th century evolutionary ideas that preceded Charles Darwin's theory of natural selection...

. This theory has come to be known as Lamarckism
Lamarckism
Lamarckism is the idea that an organism can pass on characteristics that it acquired during its lifetime to its offspring . It is named after the French biologist Jean-Baptiste Lamarck , who incorporated the action of soft inheritance into his evolutionary theories...

 and was an influence on the anti-genetic ideas of the Stalinist Soviet
Soviet Union
The Soviet Union , officially the Union of Soviet Socialist Republics , was a constitutionally socialist state that existed in Eurasia between 1922 and 1991....

 biologist Trofim Lysenko
Trofim Lysenko
Trofim Denisovich Lysenko was a Soviet agronomist of Ukrainian origin, who was director of Soviet biology under Joseph Stalin. Lysenko rejected Mendelian genetics in favor of the hybridization theories of Russian horticulturist Ivan Vladimirovich Michurin, and adopted them into a powerful...

.

Darwin's theory


In 1859, Charles Darwin set out his theory of evolution by natural selection as an explanation for adaptation and speciation. He defined natural selection as the "principle by which each slight variation [of a trait], if useful, is preserved". The concept was simple but powerful: individuals best adapted to their environments are more likely to survive and reproduce. As long as there is some variation between them, there will be an inevitable selection of individuals with the most advantageous variations. If the variations are inherited, then differential reproductive success will lead to a progressive evolution of particular populations of a species, and populations that evolve to be sufficiently different eventually become different species.

Darwin's ideas were inspired by the observations that he had made on the Beagle voyage
Second voyage of HMS Beagle
The second voyage of HMS Beagle, from 27 December 1831 to 2 October 1836, was the second survey expedition of HMS Beagle, under captain Robert FitzRoy who had taken over command of the ship on its first voyage after her previous captain committed suicide...

, and by the work of a political economist, the Reverend Thomas Malthus
Thomas Malthus
The Reverend Thomas Robert Malthus FRS was an English scholar, influential in political economy and demography. Malthus popularized the economic theory of rent....

, who in An Essay on the Principle of Population
An Essay on the Principle of Population
The book An Essay on the Principle of Population was first published anonymously in 1798 through J. Johnson . The author was soon identified as The Reverend Thomas Robert Malthus. While it was not the first book on population, it has been acknowledged as the most influential work of its era...

, noted that population (if unchecked) increases exponentially, whereas the food supply grows only arithmetically
Linear function
In mathematics, the term linear function can refer to either of two different but related concepts:* a first-degree polynomial function of one variable;* a map between two vector spaces that preserves vector addition and scalar multiplication....

; thus, inevitable limitations of resources would have demographic implications, leading to a "struggle for existence". When Darwin read Malthus in 1838 he was already primed by his work as a naturalist to appreciate the "struggle for existence" in nature and it struck him that as population outgrew resources, "favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species."

Here is Darwin's own summary of the idea, which can be found in the fourth chapter of the Origin:
If during the long course of ages and under varying conditions of life, organic beings vary at all in the several parts of their organisation, and I think this cannot be disputed; if there be, owing to the high geometrical powers of increase of each species, at some age, season, or year, a severe struggle for life, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of all organic beings to each other and to their conditions of existence, causing an infinite diversity in structure, constitution, and habits, to be advantageous to them, I think it would be a most extraordinary fact if no variation ever had occurred useful to each being's own welfare, in the same way as so many variations have occurred useful to man. But, if variations useful to any organic being do occur, assuredly individuals thus characterised will have the best chance of being preserved in the struggle for life; and from the strong principle of inheritance they will tend to produce offspring similarly characterised. This principle of preservation, I have called, for the sake of brevity, Natural Selection.


Once he had his theory "by which to work", Darwin was meticulous about gathering and refining evidence as his "prime hobby" before making his idea public. He was in the process of writing his "big book" to present his researches when the naturalist Alfred Russel Wallace
Alfred Russel Wallace
Alfred Russel Wallace, OM, FRS was a British naturalist, explorer, geographer, anthropologist and biologist...

 independently conceived of the principle and described it in an essay he sent to Darwin to forward to Charles Lyell
Charles Lyell
Sir Charles Lyell, 1st Baronet, Kt FRS was a British lawyer and the foremost geologist of his day. He is best known as the author of Principles of Geology, which popularised James Hutton's concepts of uniformitarianism – the idea that the earth was shaped by slow-moving forces still in operation...

. Lyell and Joseph Dalton Hooker
Joseph Dalton Hooker
Sir Joseph Dalton Hooker OM, GCSI, CB, MD, FRS was one of the greatest British botanists and explorers of the 19th century. Hooker was a founder of geographical botany, and Charles Darwin's closest friend...

 decided (without Wallace's knowledge) to present his essay together with unpublished writings that Darwin had sent to fellow naturalists, and On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection was read to the Linnean Society announcing co-discovery of the principle in July 1858. Darwin published a detailed account of his evidence and conclusions in On the Origin of Species in 1859. In the 3rd edition of 1861 Darwin acknowledged that others — a notable one being William Charles Wells
William Charles Wells
William Charles Wells MD FRS FRSEd , was a Scottish-American physician and printer. He lived a life of extraordinary variety, did some notable medical research, and made the first clear statement about natural selection. He applied the idea to the origin of different skin colours in human races,...

 in 1813, and Patrick Matthew
Patrick Matthew
Patrick Matthew was a Scottish landowner and fruit farmer. He published the principle of natural selection as a mechanism of evolution over a quarter-century earlier than Charles Darwin and Alfred Russel Wallace...

 in 1831 — had proposed similar ideas, but had neither developed them nor presented them in notable scientific publications.

Darwin thought of natural selection by analogy to how farmers select crops or livestock for breeding, which he called "artificial selection"; in his early manuscripts he referred to a Nature, which would do the selection. At the time, other mechanisms of evolution such as evolution by genetic drift were not yet explicitly formulated, and Darwin believed that selection was likely only part of the story: "I am convinced that [it] has been the main, but not exclusive means of modification." In a letter to Charles Lyell in September 1860, Darwin regretted the use of the term "Natural Selection", preferring the term "Natural Preservation".

For Darwin and his contemporaries, natural selection was in essence synonymous with evolution by natural selection. After the publication of On the Origin of Species, educated people generally accepted that evolution had occurred in some form. However, natural selection remained controversial as a mechanism, partly because it was perceived to be too weak to explain the range of observed characteristics of living organisms, and partly because even supporters of evolution balked at its "unguided" and non-progressive nature, a response that has been characterized as the single most significant impediment to the idea's acceptance.

However, some thinkers enthusiastically embraced natural selection; after reading Darwin, Herbert Spencer
Herbert Spencer
Herbert Spencer was an English philosopher, biologist, sociologist, and prominent classical liberal political theorist of the Victorian era....

 introduced the term survival of the fittest, which became a popular summary of the theory. The fifth edition of On the Origin of Species published in 1869 included Spencer's phrase as an alternative to natural selection, with credit given: "But the expression often used by Mr. Herbert Spencer, of the Survival of the Fittest, is more accurate, and is sometimes equally convenient." Although the phrase is still often used by non-biologists, modern biologists avoid it because it is tautological
Tautology (rhetoric)
Tautology is an unnecessary or unessential repetition of meaning, using different and dissimilar words that effectively say the same thing...

 if "fittest" is read to mean "functionally superior" and is applied to individuals rather than considered as an averaged quantity over populations.

Modern evolutionary synthesis



Natural selection relies crucially on the idea of heredity, but it was developed long before the basic concepts of genetics. Although the Austrian monk Gregor Mendel
Gregor Mendel
Gregor Johann Mendel was an Austrian scientist and Augustinian friar who gained posthumous fame as the founder of the new science of genetics. Mendel demonstrated that the inheritance of certain traits in pea plants follows particular patterns, now referred to as the laws of Mendelian inheritance...

, the father of modern genetics, was a contemporary of Darwin's, his work would lie in obscurity until the early 20th century. Only after the integration of Darwin's theory of evolution with a complex statistical appreciation of Gregor Mendel's 're-discovered' laws of inheritance did natural selection become generally accepted by scientists.

The work of Ronald Fisher
Ronald Fisher
Sir Ronald Aylmer Fisher FRS was an English statistician, evolutionary biologist, eugenicist and geneticist. Among other things, Fisher is well known for his contributions to statistics by creating Fisher's exact test and Fisher's equation...

 (who developed the required mathematical language and The Genetical Theory of Natural Selection
The Genetical Theory of Natural Selection
The Genetical Theory of Natural Selection is a book by R.A. Fisher first published in 1930 by Clarendon. It is one of the most important books of the modern evolutionary synthesis and is commonly cited in biology books.-Editions:...

), J.B.S. Haldane (who introduced the concept of the "cost" of natural selection), Sewall Wright
Sewall Wright
Sewall Green Wright was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. With R. A. Fisher and J.B.S. Haldane, he was a founder of theoretical population genetics. He is the discoverer of the inbreeding coefficient and of...

 (who elucidated the nature of selection and adaptation), Theodosius Dobzhansky
Theodosius Dobzhansky
Theodosius Grygorovych Dobzhansky ForMemRS was a prominent geneticist and evolutionary biologist, and a central figure in the field of evolutionary biology for his work in shaping the unifying modern evolutionary synthesis...

 (who established the idea that mutation, by creating genetic diversity, supplied the raw material for natural selection: see Genetics and the Origin of Species
Genetics and the Origin of Species
Genetics and the Origin of Species is a 1937 book by the twentieth century Ukrainian-American evolutionary biologist Theodosius Dobzhansky and one of the important books of the modern evolutionary synthesis. The book describes the Modern Synthesis of Evolution Theory, also known as Synthetic...

), William Hamilton
W. D. Hamilton
William Donald Hamilton FRS was a British evolutionary biologist, widely recognised as one of the greatest evolutionary theorists of the 20th century....

 (who conceived of kin selection), Ernst Mayr
Ernst Mayr
Ernst Walter Mayr was one of the 20th century's leading evolutionary biologists. He was also a renowned taxonomist, tropical explorer, ornithologist, historian of science, and naturalist...

 (who recognised the key importance of reproductive isolation for speciation: see Systematics and the Origin of Species
Systematics and the Origin of Species
Systematics and the Origin of Species from the Viewpoint of a Zoologist is a book written by zoologist and evolutionary biologist Ernst Mayr that was first published in 1942...

) and many others formed the modern evolutionary synthesis. This synthesis cemented natural selection as the foundation of evolutionary theory, where it remains today.

Impact of the idea


Darwin's ideas, along with those of Adam Smith
Adam Smith
Adam Smith was a Scottish social philosopher and a pioneer of political economy. One of the key figures of the Scottish Enlightenment, Smith is the author of The Theory of Moral Sentiments and An Inquiry into the Nature and Causes of the Wealth of Nations...

 and Karl Marx
Karl Marx
Karl Heinrich Marx was a German philosopher, economist, sociologist, historian, journalist, and revolutionary socialist. His ideas played a significant role in the development of social science and the socialist political movement...

, had a profound influence on 19th century thought. Perhaps the most radical claim of the theory of evolution through natural selection is that "elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner" evolved from the simplest forms of life by a few simple principles. This claim inspired some of Darwin's most ardent supporters—and provoked the most profound opposition. The radicalism of natural selection, according to Stephen Jay Gould
Stephen Jay Gould
Stephen Jay Gould was an American paleontologist, evolutionary biologist, and historian of science. He was also one of the most influential and widely read writers of popular science of his generation....

, lay in its power to "dethrone some of the deepest and most traditional comforts of Western thought". In particular, it challenged long-standing beliefs in such concepts as a special and exalted place for humans in the natural world and a benevolent creator whose intentions were reflected in nature's order and design.

In the words of the philosopher Daniel Dennett
Daniel Dennett
Daniel Clement Dennett is an American philosopher, writer and cognitive scientist whose research centers on the philosophy of mind, philosophy of science and philosophy of biology, particularly as those fields relate to evolutionary biology and cognitive science. He is currently the Co-director of...

, "Darwin's dangerous idea" of evolution by natural selection is a "universal acid," which cannot be kept restricted to any vessel or container, as it soon leaks out, working its way into ever-wider surroundings. Thus, in the last decades, the concept of natural selection has spread from evolutionary biology into virtually all disciplines, including evolutionary computation
Evolutionary computation
In computer science, evolutionary computation is a subfield of artificial intelligence that involves combinatorial optimization problems....

, quantum darwinism
Quantum darwinism
Quantum Darwinism is a theory explaining the emergence of the classical world from the quantum world as due to a process of Darwinian natural selection; where the many possible quantum states are selected against in favor of a stable pointer state. It is proposed by Wojciech Zurek and a group of...

, evolutionary economics
Evolutionary economics
Evolutionary economics is part of mainstream economics as well as heterodox school of economic thought that is inspired by evolutionary biology...

, evolutionary epistemology
Evolutionary epistemology
Evolutionary epistemology refers to two distinct topics - on the one hand, the biological evolution of cognitive mechanisms in animals and humans, and on the other hand, a theory in that knowledge itself evolves by natural selection....

, evolutionary psychology
Evolutionary psychology
Evolutionary psychology is an approach in the social and natural sciences that examines psychological traits such as memory, perception, and language from a modern evolutionary perspective. It seeks to identify which human psychological traits are evolved adaptations, that is, the functional...

, and cosmological natural selection. This unlimited applicability has been called Universal Darwinism
Universal darwinism
Universal Darwinism refers to a variety of approaches that extend the theory of Darwinism beyond its original domain of biological evolution on Earth...

.

Cell and molecular biology


In the 19th century, Wilhelm Roux
Wilhelm Roux
Wilhelm Roux was a German zoologist and pioneer of experimental embryology.Roux was born and educated in Jena, Germany where he attended university and studied under Ernst Haeckel. He also attended university in Berlin and Strasbourg and studied under Gustav Schwalbe, Friedrich Daniel von...

, a founder of modern embryology, wrote a book entitled « Der Kampf der Teile im Organismus » (The struggle of parts in the organism) in which he suggested that the development of an organism results from a Darwinian competition between the parts of the embryo, occurring at all levels, from molecules to organs. In recent years, a modern version of this theory has been proposed by Jean-Jacques Kupiec. According to this cellular Darwinism, stochasticity at the molecular level generates diversity in cell types whereas cell interactions impose a characteristic order on the developing embryo.

Social and psychological theory


The social implications of the theory of evolution by natural selection also became the source of continuing controversy. Friedrich Engels
Friedrich Engels
Friedrich Engels was a German industrialist, social scientist, author, political theorist, philosopher, and father of Marxist theory, alongside Karl Marx. In 1845 he published The Condition of the Working Class in England, based on personal observations and research...

, a German political philosopher and co-originator of the ideology of communism
Communism
Communism is a social, political and economic ideology that aims at the establishment of a classless, moneyless, revolutionary and stateless socialist society structured upon common ownership of the means of production...

, wrote in 1872 that "Darwin did not know what a bitter satire he wrote on mankind when he showed that free competition, the struggle for existence, which the economists celebrate as the highest historical achievement, is the normal state of the animal kingdom". Interpretation of natural selection as necessarily 'progressive', leading to increasing 'advances' in intelligence and civilisation, was used as a justification for colonialism
Colonialism
Colonialism is the establishment, maintenance, acquisition and expansion of colonies in one territory by people from another territory. It is a process whereby the metropole claims sovereignty over the colony and the social structure, government, and economics of the colony are changed by...

 and policies of eugenics
Eugenics
Eugenics is the "applied science or the bio-social movement which advocates the use of practices aimed at improving the genetic composition of a population", usually referring to human populations. The origins of the concept of eugenics began with certain interpretations of Mendelian inheritance,...

, as well as broader sociopolitical positions now described as Social Darwinism
Social Darwinism
Social Darwinism is a term commonly used for theories of society that emerged in England and the United States in the 1870s, seeking to apply the principles of Darwinian evolution to sociology and politics...

. Konrad Lorenz
Konrad Lorenz
Konrad Zacharias Lorenz was an Austrian zoologist, ethologist, and ornithologist. He shared the 1973 Nobel Prize with Nikolaas Tinbergen and Karl von Frisch...

 won the Nobel Prize in Physiology or Medicine
Nobel Prize in Physiology or Medicine
The Nobel Prize in Physiology or Medicine administered by the Nobel Foundation, is awarded once a year for outstanding discoveries in the field of life science and medicine. It is one of five Nobel Prizes established in 1895 by Swedish chemist Alfred Nobel, the inventor of dynamite, in his will...

 in 1973 for his analysis of animal behavior in terms of the role of natural selection (particularly group selection). However, in Germany in 1940, in writings that he subsequently disowned, he used the theory as a justification for policies of the Nazi state. He wrote "... selection for toughness, heroism, and social utility...must be accomplished by some human institution, if mankind, in default of selective factors, is not to be ruined by domestication-induced degeneracy. The racial idea as the basis of our state has already accomplished much in this respect." Others have developed ideas that human societies and culture evolve by mechanisms that are analogous to those that apply to evolution of species.

More recently, work among anthropologists and psychologists has led to the development of sociobiology
Sociobiology
Sociobiology is a field of scientific study which is based on the assumption that social behavior has resulted from evolution and attempts to explain and examine social behavior within that context. Often considered a branch of biology and sociology, it also draws from ethology, anthropology,...

 and later evolutionary psychology
Evolutionary psychology
Evolutionary psychology is an approach in the social and natural sciences that examines psychological traits such as memory, perception, and language from a modern evolutionary perspective. It seeks to identify which human psychological traits are evolved adaptations, that is, the functional...

, a field that attempts to explain features of human psychology in terms of adaptation to the ancestral environment. The most prominent such example, notably advanced in the early work of Noam Chomsky
Noam Chomsky
Avram Noam Chomsky is an American linguist, philosopher, cognitive scientist, and activist. He is an Institute Professor and Professor in the Department of Linguistics & Philosophy at MIT, where he has worked for over 50 years. Chomsky has been described as the "father of modern linguistics" and...

 and later by Steven Pinker
Steven Pinker
Steven Arthur Pinker is a Canadian-American experimental psychologist, cognitive scientist, linguist and popular science author...

, is the hypothesis that the human brain is adapted to acquire
Language acquisition
Language acquisition is the process by which humans acquire the capacity to perceive, produce and use words to understand and communicate. This capacity involves the picking up of diverse capacities including syntax, phonetics, and an extensive vocabulary. This language might be vocal as with...

 the grammatical
Grammar
In linguistics, grammar is the set of structural rules that govern the composition of clauses, phrases, and words in any given natural language. The term refers also to the study of such rules, and this field includes morphology, syntax, and phonology, often complemented by phonetics, semantics,...

 rules of natural language
Natural language
In the philosophy of language, a natural language is any language which arises in an unpremeditated fashion as the result of the innate facility for language possessed by the human intellect. A natural language is typically used for communication, and may be spoken, signed, or written...

. Other aspects of human behavior and social structures, from specific cultural norms such as incest avoidance to broader patterns such as gender role
Gender role
Gender roles refer to the set of social and behavioral norms that are considered to be socially appropriate for individuals of a specific sex in the context of a specific culture, which differ widely between cultures and over time...

s, have been hypothesized to have similar origins as adaptations to the early environment in which modern humans evolved. By analogy to the action of natural selection on genes, the concept of meme
Meme
A meme is "an idea, behaviour or style that spreads from person to person within a culture."A meme acts as a unit for carrying cultural ideas, symbols or practices, which can be transmitted from one mind to another through writing, speech, gestures, rituals or other imitable phenomena...

s – "units of cultural transmission", or culture's equivalents of genes undergoing selection and recombination – has arisen, first described in this form by Richard Dawkins
Richard Dawkins
Clinton Richard Dawkins, FRS, FRSL , known as Richard Dawkins, is a British ethologist, evolutionary biologist and author...

 and subsequently expanded upon by philosophers such as Daniel Dennett
Daniel Dennett
Daniel Clement Dennett is an American philosopher, writer and cognitive scientist whose research centers on the philosophy of mind, philosophy of science and philosophy of biology, particularly as those fields relate to evolutionary biology and cognitive science. He is currently the Co-director of...

 as explanations for complex cultural activities, including human consciousness
Consciousness
Consciousness is a term that refers to the relationship between the mind and the world with which it interacts. It has been defined as: subjectivity, awareness, the ability to experience or to feel, wakefulness, having a sense of selfhood, and the executive control system of the mind...

. Extensions of the theory of natural selection to such a wide range of cultural phenomena have been distinctly controversial and are not widely accepted.

Information and systems theory


In 1922, Alfred Lotka
Alfred J. Lotka
Alfred James Lotka was a US mathematician, physical chemist, and statistician, famous for his work in population dynamics and energetics. An American biophysicist best known for his proposal of the predator-prey model, developed simultaneously but independently of Vito Volterra...

 proposed that natural selection might be understood as a physical principle that could be described in terms of the use of energy
Energy
In physics, energy is an indirectly observed quantity. It is often understood as the ability a physical system has to do work on other physical systems...

 by a system, a concept that was later developed by Howard Odum
Howard T. Odum
Howard Thomas Odum was an American ecologist...

 as the maximum power principle
Maximum power principle
The maximum power principle has been proposed as the fourth principle of energetics in open system thermodynamics, where an example of an open system is a biological cell. According to Howard T...

 whereby evolutionary systems with selective advantage maximise the rate of useful energy transformation. Such concepts are sometimes relevant in the study of applied thermodynamics
Thermodynamics
Thermodynamics is a physical science that studies the effects on material bodies, and on radiation in regions of space, of transfer of heat and of work done on or by the bodies or radiation...

.

The principles of natural selection have inspired a variety of computational techniques, such as "soft" artificial life
Artificial life
Artificial life is a field of study and an associated art form which examine systems related to life, its processes, and its evolution through simulations using computer models, robotics, and biochemistry. The discipline was named by Christopher Langton, an American computer scientist, in 1986...

, that simulate selective processes and can be highly efficient in 'adapting' entities to an environment defined by a specified fitness function
Fitness function
A fitness function is a particular type of objective function that is used to summarise, as a single figure of merit, how close a given design solution is to achieving the set aims....

. For example, a class of heuristic
Heuristic
Heuristic refers to experience-based techniques for problem solving, learning, and discovery. Heuristic methods are used to speed up the process of finding a satisfactory solution, where an exhaustive search is impractical...

 optimization
Optimization (mathematics)
In mathematics, computational science, or management science, mathematical optimization refers to the selection of a best element from some set of available alternatives....

 algorithm
Algorithm
In mathematics and computer science, an algorithm is an effective method expressed as a finite list of well-defined instructions for calculating a function. Algorithms are used for calculation, data processing, and automated reasoning...

s known as genetic algorithm
Genetic algorithm
A genetic algorithm is a search heuristic that mimics the process of natural evolution. This heuristic is routinely used to generate useful solutions to optimization and search problems...

s, pioneered by John Holland
John Henry Holland
John Henry Holland is an American scientist and Professor of Psychology and Professor of Electrical Engineering and Computer Science at the University of Michigan, Ann Arbor. He is a pioneer in complex systems and nonlinear science. He is known as the father of genetic algorithms. He was awarded...

 in the 1970s and expanded upon by David E. Goldberg
David E. Goldberg
David Edward Goldberg is an American computer scientist, and professor at the department of Industrial and Enterprise Systems Engineering at the University of Illinois at Urbana-Champaign and is most noted for his seminal works in the field of genetic algorithms...

, identify optimal solutions by simulated reproduction and mutation of a population of solutions defined by an initial probability distribution
Probability distribution
In probability theory, a probability mass, probability density, or probability distribution is a function that describes the probability of a random variable taking certain values....

. Such algorithms are particularly useful when applied to problems whose solution landscape
Energy landscape
In physics, an energy landscape is a mapping of all possible conformations of a molecular entity, or the spatial positions of interacting molecules in a system, and their corresponding energy levels, typically Gibbs free energy, on a two- or three-dimensional Cartesian coordinate system.In...

 is very rough or has many local minima.

Genetic basis of natural selection


The idea of natural selection predates the understanding of genetics. We now have a much better idea of the biology underlying heritability, which is the basis of natural selection.

Genotype and phenotype



Natural selection acts on an organism's phenotype, or physical characteristics. Phenotype is determined by an organism's genetic make-up (genotype) and the environment in which the organism lives. Often, natural selection acts on specific traits of an individual, and the terms phenotype and genotype are used narrowly to indicate these specific traits.

When different organisms in a population possess different versions of a gene for a certain trait, each of these versions is known as an allele. It is this genetic variation that underlies phenotypic traits. A typical example is that certain combinations of genes for eye color
Eye color
Eye color is a polygenic phenotypic character and is determined by two distinct factors: the pigmentation of the eye's iris and the frequency-dependence of the scattering of light by the turbid medium in the stroma of the iris....

 in humans that, for instance, give rise to the phenotype of blue eyes. (On the other hand, when all the organisms in a population share the same allele for a particular trait, and this state is stable over time, the allele is said to be fixed
Fixation (population genetics)
In population genetics, fixation is the change in a gene pool from a situation where there exist at least two variants of a particular gene to a situation where only one of the alleles remains...

in that population.)

Some traits are governed by only a single gene, but most traits are influenced by the interactions of many genes. A variation in one of the many genes that contributes to a trait may have only a small effect on the phenotype; together, these genes can produce a continuum of possible phenotypic values.

Directionality of selection



When some component of a trait is heritable, selection will alter the frequencies of the different alleles, or variants of the gene that produces the variants of the trait. Selection can be divided into three classes, on the basis of its effect on allele frequencies.

Directional selection
Directional selection
In population genetics, directional selection is a mode of natural selection in which a single phenotype is favored, causing the allele frequency to continuously shift in one direction...

 occurs when a certain allele has a greater fitness than others, resulting in an increase of its frequency. This process can continue until the allele is fixed
Fixation (population genetics)
In population genetics, fixation is the change in a gene pool from a situation where there exist at least two variants of a particular gene to a situation where only one of the alleles remains...

 and the entire population shares the fitter phenotype. It is directional selection that is illustrated in the antibiotic resistance example above.

Far more common is stabilizing selection
Stabilizing selection
-Description:Stabilizing or ambidirectional selection, , is a type of natural selection in which genetic diversity decreases as the population stabilizes on a particular trait value. This is probably the most common mechanism of action for natural selection...

 (which is commonly confused with purifying selection), which lowers the frequency of alleles that have a deleterious effect on the phenotype – that is, produce organisms of lower fitness. This process can continue until the allele is eliminated from the population. Purifying selection results in functional genetic features, such as protein-coding genes
Protein biosynthesis
Protein biosynthesis is the process in which cells build or manufacture proteins. The term is sometimes used to refer only to protein translation but more often it refers to a multi-step process, beginning with amino acid synthesis and transcription of nuclear DNA into messenger RNA, which is then...

 or regulatory sequence
Regulatory sequence
A regulatory sequence is a segment of DNA where regulatory proteins such as transcription factors bind preferentially. These regulatory proteins bind to short stretches of DNA called regulatory regions, which are appropriately positioned in the genome, usually a short distance 'upstream' of the...

s, being conserved
Conserved sequence
In biology, conserved sequences are similar or identical sequences that occur within nucleic acid sequences , protein sequences, protein structures or polymeric carbohydrates across species or within different molecules produced by the same organism...

 over time due to selective pressure against deleterious variants.

Finally, a number of forms of balancing selection
Balancing selection
Balancing selection refers to a number of selective processes by which multiple alleles are actively maintained in the gene pool of a population at frequencies above that of gene mutation. This usually happens when the heterozygotes for the alleles under consideration have a higher adaptive value...

 exist, which do not result in fixation, but maintain an allele at intermediate frequencies in a population. This can occur in diploid species (that is, those that have two pairs of chromosome
Chromosome
A chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...

s) when heterozygote individuals, who have different alleles on each chromosome at a single genetic locus
Locus (genetics)
In the fields of genetics and genetic computation, a locus is the specific location of a gene or DNA sequence on a chromosome. A variant of the DNA sequence at a given locus is called an allele. The ordered list of loci known for a particular genome is called a genetic map...

, have a higher fitness than homozygote individuals that have two of the same alleles. This is called heterozygote advantage
Heterozygote advantage
A heterozygote advantage describes the case in which the heterozygote genotype has a higher relative fitness than either the homozygote dominant or homozygote recessive genotype. The specific case of heterozygote advantage is due to a single locus known as overdominance...

 or overdominance, of which the best-known example is the malaria
Malaria
Malaria is a mosquito-borne infectious disease of humans and other animals caused by eukaryotic protists of the genus Plasmodium. The disease results from the multiplication of Plasmodium parasites within red blood cells, causing symptoms that typically include fever and headache, in severe cases...

l resistance observed in heterozygous humans who carry only one copy of the gene for sickle cell anemia. Maintenance of allelic variation can also occur through disruptive or diversifying selection
Disruptive selection
Disruptive selection, also called diversifying selection, describes changes in population genetics in which extreme values for a trait are favored over intermediate values. In this case, the variance of the trait increases and the population is divided into two distinct groups...

, which favors genotypes that depart from the average in either direction (that is, the opposite of overdominance), and can result in a bimodal distribution
Bimodal distribution
In statistics, a bimodal distribution is a continuous probability distribution with two different modes. These appear as distinct peaks in the probability density function, as shown in Figure 1....

 of trait values. Finally, balancing selection can occur through frequency-dependent selection, where the fitness of one particular phenotype depends on the distribution of other phenotypes in the population. The principles of game theory
Game theory
Game theory is a mathematical method for analyzing calculated circumstances, such as in games, where a person’s success is based upon the choices of others...

 have been applied to understand the fitness distributions in these situations, particularly in the study of kin selection
Kin selection
Kin selection refers to apparent strategies in evolution that favor the reproductive success of an organism's relatives, even at a cost to the organism's own survival and reproduction. Charles Darwin was the first to discuss the concept of group/kin selection...

 and the evolution of reciprocal altruism
Reciprocal altruism
In evolutionary biology, reciprocal altruism is a behaviour whereby an organism acts in a manner that temporarily reduces its fitness while increasing another organism's fitness, with the expectation that the other organism will act in a similar manner at a later time...

.

Selection and genetic variation


A portion of all genetic variation
Genetic variation
Genetic variation, variation in alleles of genes, occurs both within and among populations. Genetic variation is important because it provides the “raw material” for natural selection. Genetic variation is brought about by mutation, a change in a chemical structure of a gene. Polyploidy is an...

 is functionally neutral in that it produces no phenotypic effect or significant difference in fitness; the hypothesis that this variation accounts for a large fraction of observed genetic diversity
Genetic diversity
Genetic diversity, the level of biodiversity, refers to the total number of genetic characteristics in the genetic makeup of a species. It is distinguished from genetic variability, which describes the tendency of genetic characteristics to vary....

 is known as the neutral theory of molecular evolution
Neutral theory of molecular evolution
The neutral theory of molecular evolution states that the vast majority of evolutionary changes at the molecular level are caused by random drift of selectively neutral mutants . The theory was introduced by Motoo Kimura in the late 1960s and early 1970s...

 and was originated by Motoo Kimura
Motoo Kimura
was a Japanese biologist best known for introducing the neutral theory of molecular evolution in 1968. He became one of the most influential theoretical population geneticists. He is remembered in genetics for his innovative use of diffusion equations to calculate the probability of fixation of...

. When genetic variation does not result in differences in fitness, selection cannot directly affect the frequency of such variation. As a result, the genetic variation at those sites will be higher than at sites where variation does influence fitness. However, after a period with no new mutation, the genetic variation at these sites will be eliminated due to genetic drift
Genetic drift
Genetic drift or allelic drift is the change in the frequency of a gene variant in a population due to random sampling.The alleles in the offspring are a sample of those in the parents, and chance has a role in determining whether a given individual survives and reproduces...

.

Mutation selection balance


Natural selection results in the reduction of genetic variation through the elimination of maladapted individuals and consequently of the mutations that caused the maladaptation. At the same time, new mutations occur, resulting in a mutation-selection balance
Mutation-selection balance
The mutation-selection balance is a classic result in population geneticsfirst derived in the 1920s by John Burdon Sanderson Haldane and R.A. Fisher.A genetic variant that is deleterious will not necessarily disappear immediately from apopulation...

. The exact outcome of the two processes depends both on the rate at which new mutations occur and on the strength of the natural selection, which is a function of how unfavorable the mutation proves to be. Consequently, changes in the mutation rate or the selection pressure will result in a different mutation-selection balance.

Genetic linkage


Genetic linkage
Genetic linkage
Genetic linkage is the tendency of certain loci or alleles to be inherited together. Genetic loci that are physically close to one another on the same chromosome tend to stay together during meiosis, and are thus genetically linked.-Background:...

 occurs when the loci
Locus (genetics)
In the fields of genetics and genetic computation, a locus is the specific location of a gene or DNA sequence on a chromosome. A variant of the DNA sequence at a given locus is called an allele. The ordered list of loci known for a particular genome is called a genetic map...

 of two alleles are linked, or in close proximity to each other on the chromosome. During the formation of gametes, recombination
Genetic recombination
Genetic recombination is a process by which a molecule of nucleic acid is broken and then joined to a different one. Recombination can occur between similar molecules of DNA, as in homologous recombination, or dissimilar molecules, as in non-homologous end joining. Recombination is a common method...

 of the genetic material results in reshuffling of the alleles. However, the chance that such a reshuffle occurs between two alleles depends on the distance between those alleles; the closer the alleles are to each other, the less likely it is that such a reshuffle will occur. Consequently, when selection targets one allele, this automatically results in selection of the other allele as well; through this mechanism, selection can have a strong influence on patterns of variation in the genome.

Selective sweep
Selective sweep
A selective sweep is the reduction or elimination of variation among the nucleotides in neighboring DNA of a mutation as the result of recent and strong positive natural selection....

s occur when an allele becomes more common in a population as a result of positive selection. As the prevalence of one allele increases, linked alleles can also become more common, whether they are neutral or even slightly deleterious. This is called genetic hitchhiking
Genetic hitchhiking
Genetic hitchhiking is the process by which an allele may increase in frequency by virtue of being linked to a gene that is positively selected. Proximity on a chromosome may allow genes to be dragged along with a selective sweep experienced by an advantageous gene nearby...

. A strong selective sweep results in a region of the genome where the positively selected haplotype
Haplotype
A haplotype in genetics is a combination of alleles at adjacent locations on the chromosome that are transmitted together...

 (the allele and its neighbors) are in essence the only ones that exist in the population.

Whether a selective sweep has occurred or not can be investigated by measuring linkage disequilibrium
Linkage disequilibrium
In population genetics, linkage disequilibrium is the non-random association of alleles at two or more loci, not necessarily on the same chromosome. It is also referred to as to as gametic phase disequilibrium , or simply gametic disequilibrium...

, or whether a given haplotype is overrepresented in the population. Normally, genetic recombination
Genetic recombination
Genetic recombination is a process by which a molecule of nucleic acid is broken and then joined to a different one. Recombination can occur between similar molecules of DNA, as in homologous recombination, or dissimilar molecules, as in non-homologous end joining. Recombination is a common method...

 results in a reshuffling of the different alleles within a haplotype, and none of the haplotypes will dominate the population. However, during a selective sweep, selection for a specific allele will also result in selection of neighboring alleles. Therefore, the presence of a block of strong linkage disequilibrium might indicate that there has been a 'recent' selective sweep near the center of the block, and this can be used to identify sites recently under selection.

Background selection is the opposite of a selective sweep. If a specific site experiences strong and persistent purifying selection, linked variation will tend to be weeded out along with it, producing a region in the genome of low overall variability. Because background selection is a result of deleterious new mutations, which can occur randomly in any haplotype, it does not produce clear blocks of linkage disequilibrium, although with low recombination it can still lead to slightly negative linkage disequilibrium overall.

See also


  • Artificial selection
    Artificial selection
    Artificial selection describes intentional breeding for certain traits, or combination of traits. The term was utilized by Charles Darwin in contrast to natural selection, in which the differential reproduction of organisms with certain traits is attributed to improved survival or reproductive...

  • Co-evolution
    Co-evolution
    In biology, coevolution is "the change of a biological object triggered by the change of a related object." Coevolution can occur at many biological levels: it can be as microscopic as correlated mutations between amino acids in a protein, or as macroscopic as covarying traits between different...

  • Dominant group (evolutionary biology)
  • Evolvability
    Evolvability
    Evolvability is defined as the capacity of a system for adaptive evolution. Evolvability is the ability of a population of organisms to not merely generate genetic diversity, but to generate adaptive genetic diversity, and thereby evolve through natural selection.In order for a biological organism...

  • Gene-centered view of evolution
    Gene-centered view of evolution
    The gene-centered view of evolution, gene selection theory or selfish gene theory holds that evolution occurs through the differential survival of competing genes, increasing the frequency of those alleles whose phenotypic effects successfully promote their own propagation, with gene defined as...

  • Genomics of domestication
    Genomics of domestication
    Genomics is the study of the structure, content, and evolution of genomes, or the entire genetic information of organisms. Domestication is the process by which humans alter the morphology and genes of targeted organisms in order to select for desirable traits.-Background:Since Domestication...

  • Negative selection
    Negative selection (natural selection)
    In natural selection, negative selection or purifying selection is the selective removal of alleles that are deleterious. This can result in stabilizing selection through the purging of deleterious variations that arise....

  • Unit of selection
    Unit of selection
    A unit of selection is a biological entity within the hierarchy of biological organisation that is subject to natural selection...


Further reading

  • For technical audiences
    • Popper, Karl
      Karl Popper
      Sir Karl Raimund Popper, CH FRS FBA was an Austro-British philosopher and a professor at the London School of Economics...

       (1978) Natural selection and the emergence of mind. Dialectica 32:339-55. See http://mertsahinoglu.com/research/karl-popper-on-the-scientific-status-of-darwins-theory-of-evolution/
    • Sober, Elliott
      Elliott Sober
      Elliott Sober is Hans Reichenbach Professor and William F. Vilas Research Professor in the Department of Philosophy at University of Wisconsin–Madison. Sober is noted for his work in philosophy of biology and general philosophy of science. Sober taught for one year at Stanford University and has...

       (1984) The Nature of Selection: Evolutionary Theory in Philosophical Focus. University of Chicago Press.
    • Williams, George C.
      George C. Williams
      Professor George Christopher Williams was an American evolutionary biologist.Williams was a professor emeritus of biology at the State University of New York at Stony Brook. He was best known for his vigorous critique of group selection. The work of Williams in this area, along with W. D...

       (1966) Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought
      Adaptation and Natural Selection
      Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought is a 1966 book by the American evolutionary biologist George C. Williams...

      .
      Oxford University Press.
    • Williams George C.
      George C. Williams
      Professor George Christopher Williams was an American evolutionary biologist.Williams was a professor emeritus of biology at the State University of New York at Stony Brook. He was best known for his vigorous critique of group selection. The work of Williams in this area, along with W. D...

       (1992) Natural Selection: Domains, Levels and Challenges. Oxford University Press.

  • For general audiences
    • Dawkins, Richard
      Richard Dawkins
      Clinton Richard Dawkins, FRS, FRSL , known as Richard Dawkins, is a British ethologist, evolutionary biologist and author...

       (1996) Climbing Mount Improbable
      Climbing Mount Improbable
      Climbing Mount Improbable is a 1996 popular science book by Richard Dawkins. The book is about probability and how it applies to the theory of evolution, and is specifically designed to debunk claims by creationists about the probability of naturalistic mechanisms like natural selection.The main...

      .
      Penguin Books, ISBN 0-670-85018-7.
    • Dennett, Daniel
      Daniel Dennett
      Daniel Clement Dennett is an American philosopher, writer and cognitive scientist whose research centers on the philosophy of mind, philosophy of science and philosophy of biology, particularly as those fields relate to evolutionary biology and cognitive science. He is currently the Co-director of...

       (1995) Darwin's Dangerous Idea: Evolution and the Meanings of Life
      Darwin's Dangerous Idea
      Darwin's Dangerous Idea: Evolution and the Meanings of Life is a book by Daniel Dennett which argues that Darwinian processes are the central organizing force that gives rise to complexity...

      .
      Simon & Schuster ISBN 0-684-82471-X.
    • Gould, Stephen Jay
      Stephen Jay Gould
      Stephen Jay Gould was an American paleontologist, evolutionary biologist, and historian of science. He was also one of the most influential and widely read writers of popular science of his generation....

       (1997) Ever Since Darwin: Reflections in Natural History. Norton, ISBN 0-393-06425-5.
    • Jones, Steve
      Steve Jones (biologist)
      John Stephen Jones is a Welsh geneticist and from 1995 to 1999 and 2008 to June 2010 was Head of the Department of Genetics, Evolution and Environment at University College London. His studies are conducted in the Galton Laboratory. He is also a television presenter and a prize-winning author on...

       (2001) Darwin's Ghost: The Origin of Species Updated. Ballantine Books ISBN 0-345-42277-5. Also published in Britain under the title Almost like a whale: the origin of species updated. Doubleday. ISBN 1-86230-025-9.
    • Lewontin, Richard
      Richard Lewontin
      Richard Charles "Dick" Lewontin is an American evolutionary biologist, geneticist and social commentator. A leader in developing the mathematical basis of population genetics and evolutionary theory, he pioneered the notion of using techniques from molecular biology such as gel electrophoresis to...

       (1978) Adaptation. Scientific American 239:212-30
    • Mayr, Ernst
      Ernst Mayr
      Ernst Walter Mayr was one of the 20th century's leading evolutionary biologists. He was also a renowned taxonomist, tropical explorer, ornithologist, historian of science, and naturalist...

       (2001) What evolution is. Weidenfeld & Nicolson, London. ISBN 0297607413
    • Weiner, Jonathan
      Jonathan Weiner
      Jonathan Weiner is a Pulitzer Prize-winning author of non-fiction books on his biology observations, in particular evolution in the Galápagos Islands, genetics, and the environment....

       (1994) The Beak of the Finch
      The Beak of the Finch
      The Beak of the Finch: A Story of Evolution in Our Time is a 1994 nonfiction book about evolutionary biology, written by Jonathan Weiner. It won the 1995 Pulitzer Prize for General Non-Fiction. The finches of the title are the Galapagos or 'Darwin's Finches,' passerine songbirds in the Galapagos...

      : A Story of Evolution in Our Time.
      Vintage Books, ISBN 0-679-73337-X.

  • Historical
    • Kohm M (2004) A Reason for Everything: Natural Selection and the English Imagination. London: Faber and Faber. ISBN 0-571-22392-3. For review, see http://human-nature.com/nibbs/05/wyhe.html van Wyhe J (2005) Human Nature Review 5:1-4

External links