Natural selection

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Modern biology began in the nineteenth century with Charles Darwin's work on evolution by natural selection.

Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Charles Darwin popularised the term "natural selection", contrasting it with artificial selection, which is intentional, whereas natural selection is not.

Variation exists within all populations of organisms. This occurs partly because random mutations arise in the genome of an individual organism, and offspring can inherit such mutations. Throughout the lives of the individuals, their genomes interact with their environments to cause variations in traits. The environment of a genome includes the molecular biology in the cell, other cells, other individuals, populations, species, as well as the abiotic environment. Because individuals with certain variants of the trait tend to survive and reproduce more than individuals with other, less successful, variants, the population evolves. Other factors affecting reproductive success include sexual selection (now often included in natural selection) and fecundity selection.

Natural selection acts on the phenotype, the characteristics of the organism which actually interact with the environment, but the genetic (heritable) basis of any phenotype that gives that phenotype a reproductive advantage may become more common in a population. Over time, this process can result in populations that specialise for particular ecological niches (microevolution) and may eventually result in speciation (the emergence of new species, macroevolution). In other words, natural selection is a key process in the evolution of a population.

Natural selection is a cornerstone of modern biology. The concept, published by Darwin and Alfred Russel Wallace in a joint presentation of papers in 1858, was elaborated in Darwin's influential 1859 book On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. He described natural selection as analogous to artificial selection, a process by which animals and plants with traits considered desirable by human breeders are systematically favoured for reproduction. The concept of natural selection originally developed in the absence of a valid theory of heredity; at the time of Darwin's writing, science had yet to develop modern theories of genetics. The union of traditional Darwinian evolution with subsequent discoveries in classical genetics formed the modern synthesis of the mid-20th century. The addition of molecular genetics has led to evolutionary developmental biology, which explains evolution at the molecular level. While genotypes can slowly change by random genetic drift, natural selection remains the primary explanation for adaptive evolution.

Historical development

Pre-Darwinian theories

Aristotle considered whether different forms could have appeared, only the useful ones surviving.

Several philosophers of the classical era, including Empedocles^[1] and his intellectual successor, the Roman poet Lucretius,^[2] 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 persist. Empedocles' idea that organisms arose entirely by the incidental workings of causes such as heat and cold was criticised by Aristotle in Book II of Physics.^[3] He posited natural teleology in its place, and believed that form was achieved for a purpose, citing the regularity of heredity in species as proof.^[4][5] Nevertheless, he accepted in his biology that new types of animals, monstrosities (τερας), can occur in very rare instances (Generation of Animals, Book IV).^[6] As quoted in Darwin's 1872 edition of The Origin of Species, Aristotle considered whether different forms (e.g., of teeth) might have appeared accidentally, but only the useful forms survived:

So what hinders the different parts [of the body] from having this merely accidental relation in nature? as the teeth, for example, grow by necessity, the front ones sharp, adapted for dividing, and the grinders flat, and serviceable for masticating the food; since they were not made for the sake of this, but it was the result of accident. And in like manner as to the other parts in which there appears to exist an adaptation to an end. Wheresoever, therefore, all things together (that is all the parts of one whole) happened like as if they were made for the sake of something, these were preserved, having been appropriately constituted by an internal spontaneity, and whatsoever things were not thus constituted, perished, and still perish.

— Aristotle, Physics, Book II, Chapter 8^[7]

But Aristotle rejected this possibility in the next paragraph, making clear that he is talking about the development of animals as embryos with the phrase "either invariably or normally come about", not the origin of species:

... Yet it is impossible that this should be the true view. For teeth and all other natural things either invariably or normally come about in a given way; but of not one of the results of chance or spontaneity is this true. We do not ascribe to chance or mere coincidence the frequency of rain in winter, but frequent rain in summer we do; nor heat in the dog-days, but only if we have it in winter. If then, it is agreed that things are either the result of coincidence or for an end, and these cannot be the result of coincidence or spontaneity, it follows that they must be for an end; and that such things are all due to nature even the champions of the theory which is before us would agree. Therefore action for an end is present in things which come to be and are by nature.

— Aristotle, Physics, Book II, Chapter 8^[8]

The struggle for existence was later described by the Islamic writer Al-Jahiz in the 9th century.^[9][10][11]

The classical arguments were reintroduced in the 18th century by Pierre Louis Maupertuis^[12] and others, including Darwin's grandfather, Erasmus 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 ideals (or typus) of created kinds. However, the theory of uniformitarianism in geology 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.^[13]

The early 19th-century zoologist 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.^[14] This theory, Lamarckism, was an influence on the Soviet biologist Trofim Lysenko's antagonism to mainstream genetic theory as late as the mid 20th century.^[15]
Between 1835 and 1837, the zoologist Edward Blyth worked on the area of variation, artificial selection, and how a similar process occurs in nature. Darwin acknowledged Blyth's ideas in the first chapter on variation of On the Origin of Species.^[16]

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".^[17] 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 and that variation is heritable, there will be an inevitable selection of individuals with the most advantageous variations. If the variations are heritable, then differential reproductive success leads to a progressive evolution of particular populations of a species, and populations that evolve to be sufficiently different eventually become different species.^[18][19]

Part of Thomas Malthus's table of population growth in England 1780–1810, from his Essay on the Principle of Population, 6th edition, 1826

Darwin's ideas were inspired by the observations that he had made on the second voyage of HMS Beagle (1831–1836), and by the work of a political economist, Thomas Robert Malthus, who, in An Essay on the Principle of Population (1798), noted that population (if unchecked) increases exponentially, whereas the food supply grows only arithmetically; thus, inevitable limitations of resources would have demographic implications, leading to a "struggle for existence".^[20] When Darwin read Malthus in 1838 he was already primed by his work as a naturalist to appreciate the "struggle for existence" in nature. 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."^[21] Darwin wrote:

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.

— Darwin summarising natural selection in the fourth chapter of On the Origin of Species^[22]

Once he had his theory, Darwin was meticulous about gathering and refining evidence before making his idea public. He was in the process of writing his "big book" to present his research when the naturalist Alfred Russel Wallace independently conceived of the principle and described it in an essay he sent to Darwin to forward to Charles Lyell. Lyell and Joseph Dalton Hooker decided 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 of London announcing co-discovery of the principle in July 1858.^[23] 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—like William Charles Wells in 1813, and Patrick Matthew in 1831—had proposed similar ideas, but had neither developed them nor presented them in notable scientific publications.^[24]

Charles Darwin noted that pigeon fanciers had created many kinds of pigeon, such as Tumblers (1, 12), Fantails (13), and Pouters (14) by selective breeding.

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 Natural Selection has been the main but not exclusive means of modification."^[25] In a letter to Charles Lyell in September 1860, Darwin regretted the use of the term "Natural Selection", preferring the term "Natural Preservation".^[26]

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,^[27] 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,^[28] a response that has been characterised as the single most significant impediment to the idea's acceptance.^[29] However, some thinkers enthusiastically embraced natural selection; after reading Darwin, Herbert Spencer introduced the phrase survival of the fittest, which became a popular summary of the theory.^[30][31] 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."^[32] Although the phrase is still often used by non-biologists, modern biologists avoid it because it is tautological if "fittest" is read to mean "functionally superior" and is applied to individuals rather than considered as an averaged quantity over populations.^[33]

The modern synthesis

Natural selection relies crucially on the idea of heredity, but developed before the basic concepts of genetics. Although the Moravian monk Gregor Mendel, the father of modern genetics, was a contemporary of Darwin's, his work lay in obscurity, only being rediscovered in 1900.^[34] With the early 20th century integration of evolution with Mendel's laws of inheritance, the so-called modern synthesis, scientists generally came to accept natural selection.^[35][36] The synthesis grew from advances in different fields. Ronald Fisher developed the required mathematical language and wrote The Genetical Theory of Natural Selection (1930).^[37] J. B. S. Haldane introduced the concept of the "cost" of natural selection.^[38][39] Sewall Wright elucidated the nature of selection and adaptation.^[40] In his book Genetics and the Origin of Species (1937), Theodosius Dobzhansky established the idea that mutation, once seen as a rival to selection, actually supplied the raw material for natural selection by creating genetic diversity.^[41][42]

A second synthesis

Evolutionary developmental biology relates the evolution of form to the precise pattern of gene activity, here gap genes in the fruit fly, during embryonic development.^[43]

Ernst Mayr recognised the key importance of reproductive isolation for speciation in his Systematics and the Origin of Species (1942).^[44] W. D. Hamilton conceived of kin selection in 1964.^[45][46] This synthesis cemented natural selection as the foundation of evolutionary theory, where it remains today. A second synthesis was brought about at the end of the 20th century by advances in molecular genetics, creating the field of evolutionary developmental biology ("evo-devo"), which seeks to explain the evolution of form in terms of the genetic regulatory programs which control the development of the embryo at molecular level. Natural selection is here understood to act on embryonic development to change the morphology of the adult body.^{[47][48][49][50]}

Terminology

The term natural selection is most often defined to operate on heritable traits, because these directly participate in evolution. However, natural selection is "blind" in the sense that changes in phenotype can give a reproductive advantage regardless of whether or not the trait is heritable. Following Darwin's primary usage, the term is used to refer both to the evolutionary consequence of blind selection and to its mechanisms.^{[27][37][51][52]} It is sometimes helpful to explicitly distinguish between selection's mechanisms and its effects; when this distinction is important, scientists define "(phenotypic) 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.^[53][54][55] Traits that cause greater reproductive success of an organism are said to be selected for, while those that reduce success are selected against.^[56]

Mechanism

Heritable variation, differential reproduction

During the industrial revolution, pollution killed many lichens, leaving tree trunks dark. A dark (melanic) morph of the peppered moth largely replaced the formerly usual light morph (both shown here). Since the moths are subject to predation by birds hunting by sight, the colour change offers better camouflage against the changed background, suggesting natural selection at work.

Natural variation occurs among the individuals of any population of organisms. Some differences may improve an individual's chances of surviving and reproducing such that its lifetime reproductive rate is increased, which means that it leaves more offspring. If the traits that give these individuals a reproductive advantage are also heritable, that is, passed from parent to offspring, then there will be differential reproduction, that is, a slightly higher proportion of fast rabbits or efficient algae in the next generation. Even if the reproductive advantage is very slight, over many generations any advantageous heritable trait becomes dominant in the population. In this way the natural environment of an organism "selects for" traits that confer a reproductive advantage, causing evolutionary change, as Darwin described.^[57] This gives the appearance of purpose, but in natural selection there is no intentional choice. Artificial selection is purposive where natural selection is not, though biologists often use teleological language to describe it.^[58]

The peppered moth exists in both light and dark colours in Great Britain, but during the industrial revolution, many of the trees on which the moths rested became blackened by soot, giving the dark-coloured moths an advantage in hiding from predators. This gave dark-coloured moths a better chance of surviving to produce dark-coloured offspring, and in just fifty years from the first dark moth being caught, nearly all of the moths in industrial Manchester were dark. The balance was reversed by the effect of the Clean Air Act 1956, and the dark moths became rare again, demonstrating the influence of natural selection on peppered moth evolution.^[59]

Fitness

The concept of fitness 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", but the precise meaning of the term is much more subtle. 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 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.^[60]

Competition

In biology, competition is an interaction between organisms in which the fitness of one is lowered by the presence of another. This may be because both rely on a limited supply of a resource such as food, water, or territory.^[61] Competition may be within or between species, and may be direct or indirect.^[62] Species less suited to compete should in theory either adapt or die out, since competition plays a powerful role in natural selection, but according to the "room to roam" theory it may be less important than expansion among larger clades.^[62][63]
Competition is modelled by r/K selection theory, which is based on Robert MacArthur and E. O. Wilson's work on island biogeography.^[64] In this theory, selective pressures drive evolution in one of two stereotyped directions: r- or K-selection.^[65] These terms, r and K, can be illustrated in a logistic model of population dynamics:^[66]

${\displaystyle {\frac {dN}{dt}}=rN\left(1-{\frac {N}{K}}\right)\qquad \!}$

where r is the growth rate of the population (N), and K is the carrying capacity of its local environmental setting. Typically, r-selected species exploit empty niches, and produce many offspring, each with a relatively low probability of surviving to adulthood. In contrast, K-selected species are strong competitors in crowded niches, and invest more heavily in much fewer offspring, each with a relatively high probability of surviving to adulthood.^[66]

Types of selection

1: directional selection: a single extreme phenotype favoured.
2, stabilizing selection: intermediate favoured over extremes.
3: disruptive selection: extremes favoured over intermediate.
X-axis: phenotypic trait
Y-axis: number of organisms
Group A: original population
Group B: after selection

Natural selection can act on any heritable phenotypic trait,^[67] and selective pressure can be produced by any aspect of the environment, including sexual selection and competition with members of the same or other species.^[68][69] 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.^[57]

Selection can be classified in several different ways, such as by its effect on a trait, on genetic diversity, by the life cycle stage where it acts, by the unit of selection, or by the resource being competed for.

Selection has different effects on traits. Stabilizing selection acts to hold a trait at a stable optimum, and in the simplest case all deviations from this optimum are selectively disadvantageous. Directional selection favours extreme values of a trait. The uncommon disruptive selection also acts during transition periods when the current mode is sub-optimal, but alters the trait in more than one direction. In particular, if the trait is quantitative and univariate then both higher and lower trait levels are favoured. Disruptive selection can be a precursor to speciation.^[57]

Alternatively, selection can be divided according to its effect on genetic diversity. Purifying or negative selection acts to remove genetic variation from the population (and is opposed by de novo mutation, which introduces new variation.^[70][71] In contrast, balancing selection acts to maintain genetic variation in a population, even in the absence of de novo mutation, by negative frequency-dependent selection. One mechanism for this is heterozygote advantage, where individuals with two different alleles have a selective advantage over individuals with just one allele. The polymorphism at the human ABO blood group locus has been explained in this way.^[72]

Different types of selection act at each life cycle stage of a sexually reproducing organism.^[73]

Another option is to classify selection by the life cycle stage at which it acts. Some biologists recognise just two types: viability (or survival) selection, which acts to increase an organism's probability of survival, and fecundity (or fertility or reproductive) selection, which acts to increase the rate of reproduction, given survival. Others split the life cycle into further components of selection. Thus viability and survival selection may be defined separately and respectively as acting to improve the probability of survival before and after reproductive age is reached, while fecundity selection may be split into additional sub-components including sexual selection, gametic selection, acting on gamete survival, and compatibility selection, acting on zygote formation.^[73]

Selection can also be classified by the level or unit of selection. Individual selection acts on the individual, in the sense that adaptations are "for" the benefit of the individual, and result from selection among individuals. Gene selection acts directly at the level of the gene. In kin selection and intragenomic conflict, gene-level selection provides a more apt explanation of the underlying process. Group selection, if it occurs, acts on groups of organisms, on the assumption that groups replicate and mutate in an analogous way to genes and individuals. There is an ongoing debate over the degree to which group selection occurs in nature.^[74]

Finally, selection can be classified according to the resource being competed for. Sexual selection results from competition for mates. Sexual selection typically proceeds via fecundity selection, sometimes at the expense of viability. Ecological selection is natural selection via any means other than sexual selection, such as kin selection, competition, and infanticide. Following Darwin, natural selection is sometimes defined as ecological selection, in which case sexual selection is considered a separate mechanism.^[75]

Sexual selection

The peacock's elaborate plumage is mentioned by Darwin as an example of sexual selection,^[76] and is a classic example of Fisherian runaway,^[77] driven to its conspicuous size and coloration through mate choice by females over many generations.

Sexual selection as first articulated by Darwin (using the example of the peacock's tail)^[76] refers specifically to competition for mates,^[78] which can be intrasexual, between individuals of the same sex, that is male–male competition, or intersexual, where one gender chooses mates, most often with males displaying and females choosing.^[79] However, in some species, mate choice is primarily by males, as in some fishes of the family Syngnathidae.^[80][81]

Phenotypic traits can be displayed in one sex and desired in the other sex, causing a positive feedback loop called a Fisherian runaway, for example, the extravagant plumage of some male birds such as the peacock.^[77] An alternate theory proposed by the same Ronald Fisher in 1930 is the sexy son hypothesis, that mothers want promiscuous sons to give them large numbers of grandchildren and so choose promiscuous fathers for their children. Aggression between members of the same sex is sometimes associated with very distinctive features, such as the antlers of stags, which are used in combat with other stags. More generally, intrasexual selection is often associated with sexual dimorphism, including differences in body size between males and females of a species.^[79]

Natural selection in action

Selection in action: resistance to antibiotics grows though the survival of individuals less affected by the antibiotic. Their offspring inherit the resistance.

 
Natural selection is seen in action in the development of antibiotic resistance in microorganisms. Since the discovery of penicillin in 1928, antibiotics have been used to fight bacterial diseases. The widespread misuse of antibiotics has selected for microbial resistance to antibiotics in clinical use, to the point that the methicillin-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.^[82] Response strategies typically include the use of different, stronger antibiotics; however, new strains of MRSA have recently emerged that are resistant even to these drugs.^[83] This is an evolutionary arms race, in which bacteria develop strains less susceptible to antibiotics, while medical researchers attempt to develop new antibiotics that can kill them. A similar situation occurs with pesticide resistance 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 suppressing male-killing activity by Wolbachia 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^[84][85]

Evolution by means of natural selection

X-ray of the left hand of a ten-year-old boy with polydactyly, caused by a mutant Hox gene

A prerequisite for natural selection to result in adaptive evolution, novel traits and speciation is the presence of heritable genetic variation that results in fitness differences. Genetic variation is the result of mutations, genetic recombinations and alterations in the karyotype (the number, shape, size and internal arrangement of the chromosomes). Any of these changes might have an effect that is highly advantageous or highly disadvantageous, but large effects are rare. In the past, most changes in the genetic material were considered neutral or close to neutral because they occurred in noncoding DNA or resulted in a synonymous substitution. However, many mutations in non-coding DNA have deleterious effects.^[86][87] Although both mutation rates and average fitness effects of mutations are dependent on the organism, a majority of mutations in humans are slightly deleterious.^[88]

Some mutations occur in "toolkit" or regulatory genes. Changes in these often 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 embryos. Some nonlethal regulatory mutations occur in HOX genes in humans, which can result in a cervical rib^[89] or polydactyly, an increase in the number of fingers or toes.^[90] When such mutations result in a higher fitness, natural selection favours these phenotypes and the novel trait spreads 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 becomes more variable and deteriorate over time, possibly resulting in a vestigial manifestation of the trait, also called evolutionary baggage. 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. A famous example of a vestigial structure, the eye of the blind mole-rat, is believed to retain function in photoperiod perception.^[91]

Speciation

Speciation requires a degree of reproductive isolation—that is, a reduction in gene flow. However, it is intrinsic to the concept of a species that hybrids are selected against, opposing the evolution of reproductive isolation, a problem that was recognised by Darwin. The problem does not occur in allopatric speciation with geographically separated populations, which can diverge with different sets of mutations. E. B. Poulton realized in 1903 that reproductive isolation could evolve through divergence, if each lineage acquired a different, incompatible allele of the same gene. Selection against the heterozygote would then directly create reproductive isolation, leading to the Bateson–Dobzhansky–Muller model, further elaborated by H. Allen Orr^[92] and Sergey Gavrilets.^[93] With reinforcement, however, natural selection can favor an increase in pre-zygotic isolation, influencing the process of speciation directly.^[94]

Genetic basis

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. 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 differences in phenotype. An example is the ABO blood type antigens in humans, where three alleles govern the phenotype.^[95]
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.^[96]

Directionality of selection

When some component of a trait is heritable, selection alters 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, stabilizing, and purifying selection.^[97] Directional selection occurs when an allele has a greater fitness than others, so that it increases in frequency, gaining an increasing share in the population. This process can continue until the allele is fixed and the entire population shares the fitter phenotype.^[98] Far more common is stabilizing 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 conserves functional genetic features, such as protein-coding genes or regulatory sequences, over time by selective pressure against deleterious variants.^[99]
Some forms of balancing selection do not result in fixation, but maintain an allele at intermediate frequencies in a population. This can occur in diploid species (with pairs of chromosomes) when heterozygous individuals (with just one copy of the allele) have a higher fitness than homozygous individuals (with two copies). This is called heterozygote advantage or over-dominance, of which the best-known example is the resistance to malaria in humans heterozygous for sickle-cell anaemia. Maintenance of allelic variation can also occur through disruptive or diversifying selection, which favours genotypes that depart from the average in either direction (that is, the opposite of over-dominance), and can result in a bimodal distribution 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 have been applied to understand the fitness distributions in these situations, particularly in the study of kin selection and the evolution of reciprocal altruism.^[100][101]

Selection, genetic variation, and drift

A portion of all genetic variation is functionally neutral, producing no phenotypic effect or significant difference in fitness. Motoo Kimura's neutral theory of molecular evolution by genetic drift proposes that this variation accounts for a large fraction of observed genetic diversity.^[102] Neutral events can radically reduce genetic variation through population bottlenecks.^[103] which among other things can cause the founder effect in initially small new populations.^[104] 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 is higher than at sites where variation does influence fitness.^[97] However, after a period with no new mutations, the genetic variation at these sites is eliminated due to genetic drift. Natural selection reduces genetic variation by eliminating maladapted individuals, and consequently the mutations that caused the maladaptation. At the same time, new mutations occur, resulting in a mutation–selection balance. 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 unfavourable the mutation proves to be.^[105]
Genetic linkage occurs when the loci of two alleles are in close proximity on a chromosome. During the formation of gametes, recombination reshuffles the alleles. The chance that such a reshuffle occurs between two alleles is inversely related to the distance between them. Selective sweeps occur when an allele becomes more common in a population as a result of positive selection. As the prevalence of one allele increases, closely linked alleles can also become more common by "genetic hitchhiking", whether they are neutral or even slightly deleterious. A strong selective sweep results in a region of the genome where the positively selected haplotype (the allele and its neighbours) are in essence the only ones that exist in the population. Selective sweeps can be detected by measuring linkage disequilibrium, or whether a given haplotype is overrepresented in the population. Since a selective sweep also results in selection of neighbouring alleles, the presence of a block of strong linkage disequilibrium might indicate a 'recent' selective sweep near the centre of the block.^[106]

Background selection is the opposite of a selective sweep. If a specific site experiences strong and persistent purifying selection, linked variation tends 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.^[107]

Impact

Darwin's ideas, along with those of Adam Smith and Karl Marx, had a profound influence on 19th century thought, including his radical claim 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.^[108] This inspired some of Darwin's most ardent supporters—and provoked the strongest opposition. Natural selection had the power, according to Stephen Jay Gould, to "dethrone some of the deepest and most traditional comforts of Western thought", such as the belief that humans have a special place in the world.^[109]
In the words of the philosopher Daniel Dennett, "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.^[110] Thus, in the last decades, the concept of natural selection has spread from evolutionary biology to other disciplines, including evolutionary computation, quantum Darwinism, evolutionary economics, evolutionary epistemology, evolutionary psychology, and cosmological natural selection. This unlimited applicability has been called universal Darwinism.^[111]

Origin of life

How life originated from inorganic matter remains an unresolved problem in biology. One prominent hypothesis is that life first appeared in the form of short self-replicating RNA polymers.^[112] On this view, life may have come into existence when RNA chains first experienced the basic conditions, as conceived by Charles Darwin, for natural selection to operate. These conditions are: heritability, variation of type, and competition for limited resources. The fitness of an early RNA replicator would likely have been a function of adaptive capacities that were intrinsic (i.e., determined by the nucleotide sequence) and the availability of resources.^[113][114] The three primary adaptive capacities could logically have been: (1) the capacity to replicate with moderate fidelity (giving rise to both heritability and variation of type), (2) the capacity to avoid decay, and (3) the capacity to acquire and process resources.^[113][114] These capacities would have been determined initially by the folded configurations (including those configurations with ribozyme activity) of the RNA replicators that, in turn, would have been encoded in their individual nucleotide sequences.^[115]

Cell and molecular biology

In 1881, the embryologist Wilhelm Roux published Der Kampf der Theile 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.^[116] In recent years, a modern version of this theory has been proposed by Jean-Jacques Kupiec. According to this cellular Darwinism, random variation at the molecular level generates diversity in cell types whereas cell interactions impose a characteristic order on the developing embryo.^[117]

Social and psychological theory

The social implications of the theory of evolution by natural selection also became the source of continuing controversy. Friedrich Engels, a German political philosopher and co-originator of the ideology of communism, wrote in 1872 that "Darwin did not know what a bitter satire he wrote on mankind, and especially on his countrymen, 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."^[118] Herbert Spencer and the eugenics advocate Francis Galton's interpretation of natural selection as necessarily progressive, leading to supposed advances in intelligence and civilisation, became a justification for colonialism, eugenics, and social Darwinism. For example, in 1940, Konrad Lorenz, in writings that he subsequently disowned, 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."^[119] Others have developed ideas that human societies and culture evolve by mechanisms analogous to those that apply to evolution of species.^[120]

More recently, work among anthropologists and psychologists has led to the development of sociobiology and later of evolutionary psychology, a field that attempts to explain features of human psychology in terms of adaptation to the ancestral environment. The most prominent example of evolutionary psychology, notably advanced in the early work of Noam Chomsky and later by Steven Pinker, is the hypothesis that the human brain has adapted to acquire the grammatical rules of natural language.^[121] Other aspects of human behaviour and social structures, from specific cultural norms such as incest avoidance to broader patterns such as gender roles, have been hypothesised 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 memes—"units of cultural transmission," or culture's equivalents of genes undergoing selection and recombination—has arisen, first described in this form by Richard Dawkins in 1976^[122] and subsequently expanded upon by philosophers such as Daniel Dennett as explanations for complex cultural activities, including human consciousness.^[123]

Information and systems theory

In 1922, Alfred J. Lotka proposed that natural selection might be understood as a physical principle that could be described in terms of the use of energy by a system,^[124][125] a concept later developed by Howard T. Odum as the maximum power principle in thermodynamics, whereby evolutionary systems with selective advantage maximise the rate of useful energy transformation.^[126]

The principles of natural selection have inspired a variety of computational techniques, such as "soft" artificial life, that simulate selective processes and can be highly efficient in 'adapting' entities to an environment defined by a specified fitness function.^[127] For example, a class of heuristic optimisation algorithms known as genetic algorithms, pioneered by John Henry Holland in the 1970s and expanded upon by David E. Goldberg,^[128] identify optimal solutions by simulated reproduction and mutation of a population of solutions defined by an initial probability distribution.^[129] Such algorithms are particularly useful when applied to problems whose energy landscape is very rough or has many local minima.^[130]

Superconductivity

From Wikipedia, the free encyclopedia

 

A magnet levitating above a high-temperature superconductor, cooled with liquid nitrogen. Persistent electric current flows on the surface of the superconductor, acting to exclude the magnetic field of the magnet (Faraday's law of induction). This current effectively forms an electromagnet that repels the magnet.

 

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Video of a Meissner effect in a high-temperature superconductor (black pellet) with a NdFeB magnet (metallic)

A high-temperature superconductor levitating above a magnet

Superconductivity is a phenomenon of exactly zero electrical resistance and expulsion of magnetic flux fields occurring in certain materials, called superconductors, when cooled below a characteristic critical temperature. It was discovered by Dutch physicist Heike Kamerlingh Onnes on April 8, 1911, in Leiden. Like ferromagnetism and atomic spectral lines, superconductivity is a quantum mechanical phenomenon. It is characterized by the Meissner effect, the complete ejection of magnetic field lines from the interior of the superconductor during its transitions into the superconducting state. The occurrence of the Meissner effect indicates that superconductivity cannot be understood simply as the idealization of perfect conductivity in classical physics.

The electrical resistance of a metallic conductor decreases gradually as temperature is lowered. In ordinary conductors, such as copper or silver, this decrease is limited by impurities and other defects. Even near absolute zero, a real sample of a normal conductor shows some resistance. In a superconductor, the resistance drops abruptly to zero when the material is cooled below its critical temperature. An electric current through a loop of superconducting wire can persist indefinitely with no power source.^[1][2][3][4]

In 1986, it was discovered that some cuprate-perovskite ceramic materials have a critical temperature above 90 K (−183 °C).^[5] Such a high transition temperature is theoretically impossible for a conventional superconductor, leading the materials to be termed high-temperature superconductors. The cheaply-available coolant liquid nitrogen boils at 77 K, and thus superconduction at higher temperatures than this facilitates many experiments and applications that are less practical at lower temperatures.

Classification

There are many criteria by which superconductors are classified. The most common are:

Response to a magnetic field

A superconductor can be Type I, meaning it has a single critical field, above which all superconductivity is lost and below which the magnetic field is completely expelled from the superconductor; or Type II, meaning it has two critical fields, between which it allows partial penetration of the magnetic field through isolated points. These points are called vortices. Furthermore, in multicomponent superconductors it is possible to have combination of the two behaviours. In that case the superconductor is of Type-1.5.

By theory of operation

It is conventional if it can be explained by the BCS theory or its derivatives, or unconventional, otherwise.^[6]

By critical temperature

A superconductor is generally considered high-temperature if it reaches a superconducting state when cooled using liquid nitrogen – that is, at only T_c > 77 K) – or low-temperature if more aggressive cooling techniques are required to reach its critical temperature.

By material

Superconductor material classes include chemical elements (e.g. mercury or lead), alloys (such as niobium-titanium, germanium-niobium, and niobium nitride), ceramics (YBCO and magnesium diboride), superconducting pnictides (like fluorine-doped LaOFeAs) or organic superconductors (fullerenes and carbon nanotubes; though perhaps these examples should be included among the chemical elements, as they are composed entirely of carbon).

Elementary properties of superconductors

Most of the physical properties of superconductors vary from material to material, such as the heat capacity and the critical temperature, critical field, and critical current density at which superconductivity is destroyed.

On the other hand, there is a class of properties that are independent of the underlying material. For instance, all superconductors have exactly zero resistivity to low applied currents when there is no magnetic field present or if the applied field does not exceed a critical value. The existence of these "universal" properties implies that superconductivity is a thermodynamic phase, and thus possesses certain distinguishing properties which are largely independent of microscopic details.

Zero electrical DC resistance

Electric cables for accelerators at CERN. Both the massive and slim cables are rated for 12,500 A. Top: regular cables for LEP; bottom: superconductor-based cables for the LHC

 

Cross section of a preform superconductor rod from abandoned Texas Superconducting Super Collider (SSC).

The simplest method to measure the electrical resistance of a sample of some material is to place it in an electrical circuit in series with a current source I and measure the resulting voltage V across the sample. The resistance of the sample is given by Ohm's law as R = V / I. If the voltage is zero, this means that the resistance is zero.

Superconductors are also able to maintain a current with no applied voltage whatsoever, a property exploited in superconducting electromagnets such as those found in MRI machines. Experiments have demonstrated that currents in superconducting coils can persist for years without any measurable degradation. Experimental evidence points to a current lifetime of at least 100,000 years. Theoretical estimates for the lifetime of a persistent current can exceed the estimated lifetime of the universe, depending on the wire geometry and the temperature.^[3] In practice, currents injected in superconducting coils have persisted for more than 22 years in superconducting gravimeters.^[7][8] In such instruments, the measurement principle is based on the monitoring of the levitation of a superconducting niobium sphere of mass 4 grams.

In a normal conductor, an electric current may be visualized as a fluid of electrons moving across a heavy ionic lattice. The electrons are constantly colliding with the ions in the lattice, and during each collision some of the energy carried by the current is absorbed by the lattice and converted into heat, which is essentially the vibrational kinetic energy of the lattice ions. As a result, the energy carried by the current is constantly being dissipated. This is the phenomenon of electrical resistance and Joule heating.

The situation is different in a superconductor. In a conventional superconductor, the electronic fluid cannot be resolved into individual electrons. Instead, it consists of bound pairs of electrons known as Cooper pairs. This pairing is caused by an attractive force between electrons from the exchange of phonons. Due to quantum mechanics, the energy spectrum of this Cooper pair fluid possesses an energy gap, meaning there is a minimum amount of energy ΔE that must be supplied in order to excite the fluid. Therefore, if ΔE is larger than the thermal energy of the lattice, given by kT, where k is Boltzmann's constant and T is the temperature, the fluid will not be scattered by the lattice. The Cooper pair fluid is thus a superfluid, meaning it can flow without energy dissipation.

In a class of superconductors known as type II superconductors, including all known high-temperature superconductors, an extremely low but nonzero resistivity appears at temperatures not too far below the nominal superconducting transition when an electric current is applied in conjunction with a strong magnetic field, which may be caused by the electric current. This is due to the motion of magnetic vortices in the electronic superfluid, which dissipates some of the energy carried by the current. If the current is sufficiently small, the vortices are stationary, and the resistivity vanishes. The resistance due to this effect is tiny compared with that of non-superconducting materials, but must be taken into account in sensitive experiments. However, as the temperature decreases far enough below the nominal superconducting transition, these vortices can become frozen into a disordered but stationary phase known as a "vortex glass". Below this vortex glass transition temperature, the resistance of the material becomes truly zero.

Superconducting phase transition

Behavior of heat capacity (c_v, blue) and resistivity (ρ, green) at the superconducting phase transition

In superconducting materials, the characteristics of superconductivity appear when the temperature T is lowered below a critical temperature T_c. The value of this critical temperature varies from material to material. Conventional superconductors usually have critical temperatures ranging from around 20 K to less than 1 K. Solid mercury, for example, has a critical temperature of 4.2 K. As of 2009, the highest critical temperature found for a conventional superconductor is 39 K for magnesium diboride (MgB₂),^[9][10] although this material displays enough exotic properties that there is some doubt about classifying it as a "conventional" superconductor.^[11] Cuprate superconductors can have much higher critical temperatures: YBa₂Cu₃O₇, one of the first cuprate superconductors to be discovered, has a critical temperature of 92 K, and mercury-based cuprates have been found with critical temperatures in excess of 130 K. The explanation for these high critical temperatures remains unknown. Electron pairing due to phonon exchanges explains superconductivity in conventional superconductors, but it does not explain superconductivity in the newer superconductors that have a very high critical temperature.

Similarly, at a fixed temperature below the critical temperature, superconducting materials cease to superconduct when an external magnetic field is applied which is greater than the critical magnetic field. This is because the Gibbs free energy of the superconducting phase increases quadratically with the magnetic field while the free energy of the normal phase is roughly independent of the magnetic field. If the material superconducts in the absence of a field, then the superconducting phase free energy is lower than that of the normal phase and so for some finite value of the magnetic field (proportional to the square root of the difference of the free energies at zero magnetic field) the two free energies will be equal and a phase transition to the normal phase will occur. More generally, a higher temperature and a stronger magnetic field lead to a smaller fraction of electrons that are superconducting and consequently to a longer London penetration depth of external magnetic fields and currents. The penetration depth becomes infinite at the phase transition.

The onset of superconductivity is accompanied by abrupt changes in various physical properties, which is the hallmark of a phase transition. For example, the electronic heat capacity is proportional to the temperature in the normal (non-superconducting) regime. At the superconducting transition, it suffers a discontinuous jump and thereafter ceases to be linear. At low temperatures, it varies instead as e^−α/T for some constant, α. This exponential behavior is one of the pieces of evidence for the existence of the energy gap.

The order of the superconducting phase transition was long a matter of debate. Experiments indicate that the transition is second-order, meaning there is no latent heat. However, in the presence of an external magnetic field there is latent heat, because the superconducting phase has a lower entropy below the critical temperature than the normal phase. It has been experimentally demonstrated^[12] that, as a consequence, when the magnetic field is increased beyond the critical field, the resulting phase transition leads to a decrease in the temperature of the superconducting material.

Calculations in the 1970s suggested that it may actually be weakly first-order due to the effect of long-range fluctuations in the electromagnetic field. In the 1980s it was shown theoretically with the help of a disorder field theory, in which the vortex lines of the superconductor play a major role, that the transition is of second order within the type II regime and of first order (i.e., latent heat) within the type I regime, and that the two regions are separated by a tricritical point.^[13] The results were strongly supported by Monte Carlo computer simulations.^[14]

Meissner effect

When a superconductor is placed in a weak external magnetic field H, and cooled below its transition temperature, the magnetic field is ejected. The Meissner effect does not cause the field to be completely ejected but instead the field penetrates the superconductor but only to a very small distance, characterized by a parameter λ, called the London penetration depth, decaying exponentially to zero within the bulk of the material. The Meissner effect is a defining characteristic of superconductivity. For most superconductors, the London penetration depth is on the order of 100 nm.
The Meissner effect is sometimes confused with the kind of diamagnetism one would expect in a perfect electrical conductor: according to Lenz's law, when a changing magnetic field is applied to a conductor, it will induce an electric current in the conductor that creates an opposing magnetic field. In a perfect conductor, an arbitrarily large current can be induced, and the resulting magnetic field exactly cancels the applied field.

The Meissner effect is distinct from this—it is the spontaneous expulsion which occurs during transition to superconductivity. Suppose we have a material in its normal state, containing a constant internal magnetic field. When the material is cooled below the critical temperature, we would observe the abrupt expulsion of the internal magnetic field, which we would not expect based on Lenz's law.

The Meissner effect was given a phenomenological explanation by the brothers Fritz and Heinz London, who showed that the electromagnetic free energy in a superconductor is minimized provided

${\displaystyle \nabla ^{2}\mathbf {H} =\lambda ^{-2}\mathbf {H} \,}$

where H is the magnetic field and λ is the London penetration depth.

This equation, which is known as the London equation, predicts that the magnetic field in a superconductor decays exponentially from whatever value it possesses at the surface.

A superconductor with little or no magnetic field within it is said to be in the Meissner state. The Meissner state breaks down when the applied magnetic field is too large. Superconductors can be divided into two classes according to how this breakdown occurs. In Type I superconductors, superconductivity is abruptly destroyed when the strength of the applied field rises above a critical value H_c. Depending on the geometry of the sample, one may obtain an intermediate state^[15] consisting of a baroque pattern^[16] of regions of normal material carrying a magnetic field mixed with regions of superconducting material containing no field. In Type II superconductors, raising the applied field past a critical value H_c1 leads to a mixed state (also known as the vortex state) in which an increasing amount of magnetic flux penetrates the material, but there remains no resistance to the flow of electric current as long as the current is not too large. At a second critical field strength H_c2, superconductivity is destroyed. The mixed state is actually caused by vortices in the electronic superfluid, sometimes called fluxons because the flux carried by these vortices is quantized. Most pure elemental superconductors, except niobium and carbon nanotubes, are Type I, while almost all impure and compound superconductors are Type II.

London moment

Conversely, a spinning superconductor generates a magnetic field, precisely aligned with the spin axis. The effect, the London moment, was put to good use in Gravity Probe B. This experiment measured the magnetic fields of four superconducting gyroscopes to determine their spin axes. This was critical to the experiment since it is one of the few ways to accurately determine the spin axis of an otherwise featureless sphere.

History of superconductivity

Heike Kamerlingh Onnes (right), the discoverer of superconductivity. Paul Ehrenfest, Hendrik Lorentz, Niels Bohr stand to his left.

Superconductivity was discovered on April 8, 1911 by Heike Kamerlingh Onnes, who was studying the resistance of solid mercury at cryogenic temperatures using the recently produced liquid helium as a refrigerant. At the temperature of 4.2 K, he observed that the resistance abruptly disappeared.^[17] In the same experiment, he also observed the superfluid transition of helium at 2.2 K, without recognizing its significance. The precise date and circumstances of the discovery were only reconstructed a century later, when Onnes's notebook was found.^[18] In subsequent decades, superconductivity was observed in several other materials. In 1913, lead was found to superconduct at 7 K, and in 1941 niobium nitride was found to superconduct at 16 K.

Great efforts have been devoted to finding out how and why superconductivity works; the important step occurred in 1933, when Meissner and Ochsenfeld discovered that superconductors expelled applied magnetic fields, a phenomenon which has come to be known as the Meissner effect.^[19] In 1935, Fritz and Heinz London showed that the Meissner effect was a consequence of the minimization of the electromagnetic free energy carried by superconducting current.^[20]

London theory

The first phenomenological theory of superconductivity was London theory. It was put forward by the brothers Fritz and Heinz London in 1935, shortly after the discovery that magnetic fields are expelled from superconductors. A major triumph of the equations of this theory is their ability to explain the Meissner effect,^[19] wherein a material exponentially expels all internal magnetic fields as it crosses the superconducting threshold. By using the London equation, one can obtain the dependence of the magnetic field inside the superconductor on the distance to the surface.^[21]

There are two London equations:

${\displaystyle {\frac {\partial \mathbf {j} _{s}}{\partial t}}={\frac {n_{s}e^{2}}{m}}\mathbf {E} ,\qquad \mathbf {\nabla } \times \mathbf {j} _{s}=-{\frac {n_{s}e^{2}}{m}}\mathbf {B} .}$

The first equation follows from Newton's second law for superconducting electrons.

Conventional theories (1950s)

During the 1950s, theoretical condensed matter physicists arrived at an understanding of "conventional" superconductivity, through a pair of remarkable and important theories: the phenomenological Ginzburg-Landau theory (1950) and the microscopic BCS theory (1957).^[22][23]
In 1950, the phenomenological Ginzburg-Landau theory of superconductivity was devised by Landau and Ginzburg.^[24] This theory, which combined Landau's theory of second-order phase transitions with a Schrödinger-like wave equation, had great success in explaining the macroscopic properties of superconductors. In particular, Abrikosov showed that Ginzburg-Landau theory predicts the division of superconductors into the two categories now referred to as Type I and Type II. Abrikosov and Ginzburg were awarded the 2003 Nobel Prize for their work (Landau had received the 1962 Nobel Prize for other work, and died in 1968). The four-dimensional extension of the Ginzburg-Landau theory, the Coleman-Weinberg model, is important in quantum field theory and cosmology.

Also in 1950, Maxwell and Reynolds et al. found that the critical temperature of a superconductor depends on the isotopic mass of the constituent element.^[25][26] This important discovery pointed to the electron-phonon interaction as the microscopic mechanism responsible for superconductivity.

The complete microscopic theory of superconductivity was finally proposed in 1957 by Bardeen, Cooper and Schrieffer.^[23] This BCS theory explained the superconducting current as a superfluid of Cooper pairs, pairs of electrons interacting through the exchange of phonons. For this work, the authors were awarded the Nobel Prize in 1972.

The BCS theory was set on a firmer footing in 1958, when N. N. Bogolyubov showed that the BCS wavefunction, which had originally been derived from a variational argument, could be obtained using a canonical transformation of the electronic Hamiltonian.^[27] In 1959, Lev Gor'kov showed that the BCS theory reduced to the Ginzburg-Landau theory close to the critical temperature.^[28][29]

Generalizations of BCS theory for conventional superconductors form the basis for understanding of the phenomenon of superfluidity, because they fall into the lambda transition universality class. The extent to which such generalizations can be applied to unconventional superconductors is still controversial.

Further history

The first practical application of superconductivity was developed in 1954 with Dudley Allen Buck's invention of the cryotron.^[30] Two superconductors with greatly different values of critical magnetic field are combined to produce a fast, simple switch for computer elements.

Soon after discovering superconductivity in 1911, Kamerlingh Onnes attempted to make an electromagnet with superconducting windings but found that relatively low magnetic fields destroyed superconductivity in the materials he investigated. Much later, in 1955, G.B. Yntema ^[31] succeeded in constructing a small 0.7-tesla iron-core electromagnet with superconducting niobium wire windings. Then, in 1961, J.E. Kunzler, E. Buehler, F.S.L. Hsu, and J.H. Wernick ^[32] made the startling discovery that, at 4.2 kelvin, a compound consisting of three parts niobium and one part tin, was capable of supporting a current density of more than 100,000 amperes per square centimeter in a magnetic field of 8.8 tesla. Despite being brittle and difficult to fabricate, niobium-tin has since proved extremely useful in supermagnets generating magnetic fields as high as 20 tesla. In 1962 T.G. Berlincourt and R.R. Hake ^[33][34] discovered that alloys of niobium and titanium are suitable for applications up to 10 tesla. Promptly thereafter, commercial production of niobium-titanium supermagnet wire commenced at Westinghouse Electric Corporation and at Wah Chang Corporation. Although niobium-titanium boasts less-impressive superconducting properties than those of niobium-tin, niobium-titanium has, nevertheless, become the most widely used “workhorse” supermagnet material, in large measure a consequence of its very-high ductility and ease of fabrication. However, both niobium-tin and niobium-titanium find wide application in MRI medical imagers, bending and focusing magnets for enormous high-energy-particle accelerators, and a host of other applications. Conectus, a European superconductivity consortium, estimated that in 2014, global economic activity for which superconductivity was indispensable amounted to about five billion euros, with MRI systems accounting for about 80% of that total.

In 1962, Josephson made the important theoretical prediction that a supercurrent can flow between two pieces of superconductor separated by a thin layer of insulator.^[35] This phenomenon, now called the Josephson effect, is exploited by superconducting devices such as SQUIDs. It is used in the most accurate available measurements of the magnetic flux quantum Φ₀ = h/(2e), where h is the Planck constant. Coupled with the quantum Hall resistivity, this leads to a precise measurement of the Planck constant. Josephson was awarded the Nobel Prize for this work in 1973.

In 2008, it was proposed that the same mechanism that produces superconductivity could produce a superinsulator state in some materials, with almost infinite electrical resistance.^[36]

High-temperature superconductivity

Timeline of superconducting materials

Until 1986, physicists had believed that BCS theory forbade superconductivity at temperatures above about 30 K. In that year, Bednorz and Müller discovered superconductivity in a lanthanum-based cuprate perovskite material, which had a transition temperature of 35 K (Nobel Prize in Physics, 1987).^[5] It was soon found that replacing the lanthanum with yttrium (i.e., making YBCO) raised the critical temperature to 92 K.^[37]

This temperature jump is particularly significant, since it allows liquid nitrogen as a refrigerant, replacing liquid helium.^[37] This can be important commercially because liquid nitrogen can be produced relatively cheaply, even on-site. Also, the higher temperatures help avoid some of the problems that arise at liquid helium temperatures, such as the formation of plugs of frozen air that can block cryogenic lines and cause unanticipated and potentially hazardous pressure buildup.^[38][39]

Many other cuprate superconductors have since been discovered, and the theory of superconductivity in these materials is one of the major outstanding challenges of theoretical condensed matter physics.^[40] There are currently two main hypotheses – the resonating-valence-bond theory, and spin fluctuation which has the most support in the research community.^[41] The second hypothesis proposed that electron pairing in high-temperature superconductors is mediated by short-range spin waves known as paramagnons.^{[42][43][dubious – discuss]}

Since about 1993, the highest-temperature superconductor has been a ceramic material consisting of mercury, barium, calcium, copper and oxygen (HgBa₂Ca₂Cu₃O_8+δ) with T_c = 133–138 K.^[44][45] The latter experiment (138 K) still awaits experimental confirmation, however.

In February 2008, an iron-based family of high-temperature superconductors was discovered.^[46][47] Hideo Hosono, of the Tokyo Institute of Technology, and colleagues found lanthanum oxygen fluorine iron arsenide (LaO_1−xF_xFeAs), an oxypnictide that superconducts below 26 K. Replacing the lanthanum in LaO_1−xF_xFeAs with samarium leads to superconductors that work at 55 K.^[48]

In May 2014, hydrogen sulfide (H
₂S) was predicted to be a high-temperature superconductor with a transition temperature of 80 K at 160 gigapascals of pressure.^[49] In 2015, H
₂S has been observed to exhibit superconductivity at below 203 K but at extremely high pressures — around 150 gigapascals.^[50]

Applications

 
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Video of superconducting levitation of YBCO

Superconducting magnets are some of the most powerful electromagnets known. They are used in MRI/NMR machines, mass spectrometers, the beam-steering magnets used in particle accelerators and plasma confining magnets in some tokamaks. They can also be used for magnetic separation, where weakly magnetic particles are extracted from a background of less or non-magnetic particles, as in the pigment industries.

In the 1950s and 1960s, superconductors were used to build experimental digital computers using cryotron switches. More recently, superconductors have been used to make digital circuits based on rapid single flux quantum technology and RF and microwave filters for mobile phone base stations.

Superconductors are used to build Josephson junctions which are the building blocks of SQUIDs (superconducting quantum interference devices), the most sensitive magnetometers known. SQUIDs are used in scanning SQUID microscopes and magnetoencephalography. Series of Josephson devices are used to realize the SI volt. Depending on the particular mode of operation, a superconductor-insulator-superconductor Josephson junction can be used as a photon detector or as a mixer. The large resistance change at the transition from the normal- to the superconducting state is used to build thermometers in cryogenic micro-calorimeter photon detectors. The same effect is used in ultrasensitive bolometers made from superconducting materials.

Other early markets are arising where the relative efficiency, size and weight advantages of devices based on high-temperature superconductivity outweigh the additional costs involved. For example, in wind turbines the lower weight and volume of superconducting generators could lead to savings in construction and tower costs, offsetting the higher costs for the generator and lowering the total LCOE.^[51]

Promising future applications include high-performance smart grid, electric power transmission, transformers, power storage devices, electric motors (e.g. for vehicle propulsion, as in vactrains or maglev trains), magnetic levitation devices, fault current limiters, enhancing spintronic devices with superconducting materials,^[52] and superconducting magnetic refrigeration. However, superconductivity is sensitive to moving magnetic fields so applications that use alternating current (e.g. transformers) will be more difficult to develop than those that rely upon direct current. Compared to traditional power lines superconducting transmission lines are more efficient and require only a fraction of the space, which would not only lead to a better environmental performance but could also improve public acceptance for expansion of the electric grid.^[53]

Nobel Prizes for superconductivity

• Heike Kamerlingh Onnes (1913), "for his investigations on the properties of matter at low temperatures which led, inter alia, to the production of liquid helium"
• John Bardeen, Leon N. Cooper, and J. Robert Schrieffer (1972), "for their jointly developed theory of superconductivity, usually called the BCS-theory"
• Leo Esaki, Ivar Giaever, and Brian D. Josephson (1973), "for their experimental discoveries regarding tunneling phenomena in semiconductors and superconductors, respectively," and "for his theoretical predictions of the properties of a supercurrent through a tunnel barrier, in particular those phenomena which are generally known as the Josephson effects"
• Georg Bednorz and K. Alex Müller (1987), "for their important break-through in the discovery of superconductivity in ceramic materials"
• Alexei A. Abrikosov, Vitaly L. Ginzburg, and Anthony J. Leggett (2003), "for pioneering contributions to the theory of superconductors and superfluids"^[54]
• Michael Kosterlitz, Duncan Haldane, David J. Thouless (2016)