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Peppered moth evolution

The evolution of the peppered moth is an evolutionary instance of directional colour change in the moth population as a consequence of air pollution during the Industrial Revolution. The frequency of dark-coloured moths increased at that time, an example of industrial melanism. Later, when pollution was reduced, the light-coloured form again predominated. Industrial melanism in the peppered moth was an early test of Charles Darwin's natural selection in action, and it remains a classic example in the teaching of evolution.[1][2][3] In 1978, Sewall Wright described it as "the clearest case in which a conspicuous evolutionary process has actually been observed."[4][5]

Biston betularia f. typica, the white-bodied peppered moth.
Biston betularia f. carbonaria, the black-bodied peppered moth.

The dark-coloured or melanic form of the peppered moth (var. carbonaria) was rare, though a specimen had been collected by 1811. After field collection in 1848 from Manchester, an industrial city in England, the frequency of the variety was found to have increased drastically. By the end of the 19th century it almost completely outnumbered the original light-coloured type (var. typica), with a record of 98% in 1895.[6] The evolutionary importance of the moth was only speculated upon during Darwin's lifetime. It was 14 years after Darwin's death, in 1896, that J. W. Tutt presented it as a case of natural selection.[7] Because of this, the idea spread widely, and more people came to believe in Darwin's theory.

Bernard Kettlewell was the first to investigate the evolutionary mechanism behind peppered moth adaptation, between 1953 and 1956. He found that a light-coloured body was an effective camouflage in a clean environment, such as in rural Dorset, while the dark colour was beneficial in a polluted environment like industrial Birmingham. This selective survival was due to birds, which easily caught dark moths on clean trees and white moths on trees darkened with soot. The story, supported by Kettlewell's experiment, became the canonical example of Darwinian evolution and evidence for natural selection used in standard textbooks.[8][9][10]

However, failure to replicate the experiment and Theodore David Sargent's criticism of Kettlewell's methods in the late 1960s led to general skepticism. When Judith Hooper's Of Moths and Men was published in 2002, Kettlewell's story was more sternly attacked, and accused of fraud. The criticism became a major argument for creationists. Michael Majerus was the principal defender. His seven-year experiment beginning in 2001, the most elaborate of its kind in population biology, the results of which were published posthumously in 2012, vindicated Kettlewell's work in great detail. This restored peppered moth evolution as "the most direct evidence", and "one of the clearest and most easily understood examples of Darwinian evolution in action".[11]

Origin and evolution

 
Typica and carbonaria morphs on the same tree. The light-coloured typica (below the bark's scar) is nearly invisible on this pollution-free tree, camouflaging it from predators.

Before the Industrial Revolution, the black form of the peppered moth was rare. The first black specimen (of unknown origin) was collected before 1811, and kept in the University of Oxford.[12][13][14] The first live specimen was caught by R. S. Edleston in Manchester, England in 1848, but he reported this only 16 years later in 1864, in The Entomologist.[15] Edleston notes that by 1864 it was the more common type of moth in his garden in Manchester. The light-bodied moths were able to blend in with the light-coloured lichens and tree bark, and the less common black moths were more likely to be eaten by birds. As a result of the common light-coloured lichens and English trees, therefore, the light-coloured moths were much more effective at hiding from predators, and the frequency of the dark allele was very low, at about 0.01%.[16]

During the early decades of the Industrial Revolution in England, the countryside between London and Manchester became blanketed with soot from the new coal-burning factories. Many of the light-bodied lichens died from sulphur dioxide emissions, and the trees became darkened. This led to an increase in bird predation for light-coloured moths, as they no longer blended in as well in their polluted ecosystem: indeed, their bodies now dramatically contrasted with the colour of the bark. Dark-coloured moths, on the other hand, were camouflaged very well by the blackened trees.[17] The population of dark-coloured moth rapidly increased. By the mid-19th century, the number of dark-coloured moths had risen noticeably, and by 1895, the percentage of dark-coloured moths in Manchester was reported at 98%, a dramatic change (of almost 100%) from the original frequency.[17] This effect of industrialization in body colour led to the coining of the term "industrial melanism".[3]

The implication that industrial melanism could be evidence supporting Charles Darwin's theory of natural selection was noticed during his lifetime. Albert Brydges Farn (1841–1921), a British entomologist, wrote to Darwin on 18 November 1878 to discuss his observation of colour variations in the Annulet moth (then Gnophos obscurata, now Charissa obscurata). He noted the existence of dark moths in peat in the New Forest, brown moths on clay and red soil in Herefordshire, and white moths on chalk cliffs in Lewes, and suggested that this variation was an example of "survival of the fittest". He told Darwin that he had found dark moths on a chalk slope where the foliage had been blackened by smoke from lime kilns, and he had also heard that white moths had become less common at Lewes after lime kilns had been in operation for a few years.[18] Darwin does not seem to have responded to this information, possibly because he thought natural selection would be a much slower process.[19] A scientific explanation of moth coloration was only published in 1896, 14 years after Darwin's death, when J. W. Tutt explicitly linked peppered moth melanism to natural selection.[16]

Rise and fall of phenotype frequency

Melanism has appeared in both European and North American peppered moth populations. Information about the rise in frequency is scarce. Much more is known about the subsequent fall in phenotype frequency, as it has been measured by lepidopterists using moth traps.

Steward compiled data for the first recordings of the peppered moth by locality, and deduced that the carbonaria morph was the result of a single mutation that subsequently spread. By 1895, it had reached a reported frequency of 98% in Manchester.[20]

From around 1962 to the present, the phenotype frequency of carbonaria has steadily fallen in line with cleaner air around industrial cities. Its decline has been measured more accurately than its rise, through more rigorous scientific studies. Notably, Kettlewell conducted a national survey in 1956, Bruce Grant conducted a similar one in early 1996,[21] and L.M. Cook in 2003.[22]

Similar results were found in North America. Melanic forms have not been found in Japan. It is believed that this is because peppered moths in Japan do not inhabit industrialised regions.[22]

Genetics

Tutt was the first to propose the "differential bird predation hypothesis" in 1896, as a mechanism of natural selection. The melanic morphs were better camouflaged against the bark of trees without foliose lichen, whereas the typica morphs were better camouflaged against trees with lichens. As a result, birds would find and eat those morphs that were not camouflaged with increased frequency.[23]

In 1924, J.B.S. Haldane calculated, using a simple general selection model, the selective advantage necessary for the recorded natural evolution of peppered moths, based on the assumption that in 1848 the frequency of dark-coloured moths was 2%, and by 1895 it was 95%. The dark-coloured, or melanic, form would have had to be 50% more fit than the typical, light-coloured form. Even taking into consideration possible errors in the model, this reasonably excluded the stochastic process of genetic drift, because the changes were too fast.[24] Haldane's statistical analysis of selection for the melanic variant in peppered moths became a well known part of his effort to demonstrate that mathematical models that combined natural selection with Mendelian genetics could explain evolution – an effort that played a key role in the foundation of the discipline of population genetics, and the beginnings of the modern synthesis of evolutionary theory with genetics.[25]

The peppered moth Biston betularia is also a model of parallel evolution in the incidence of melanism in the British form (f. carbonaria) and the American form (f. swettaria) as they are indistinguishable in appearance. Genetic analysis indicates that both phenotypes are inherited as autosomal dominants. Cross hybridizations indicate that the phenotypes are produced by alleles at a single locus.[26]

The gene for carbonaria in B. betularia was thought to be in a region of chromosome 17. It was later concluded that the gene could not be in that region, because none of the genes in the chromosome coded for either wing pattern or melanisation. The region that was used to find it was the first intron of the orthologue of the cortex gene in Drosophila. Through elimination of candidates within the region based on rarity, a 21,925 base pair insert remained. The insert, labelled carb-TE, is a class II transposable element that has an approximately 9-kb non-repetitive sequence tandemly repeated two and one third times. There are 6 base pairs of inverted repeats and duplicated 4 base pairs at the target site not present in typica moths. Carb-TE has higher expression during the stage of rapid wing disc morphogenesis. The mechanism of how the gene increases expression, and whether it is the only gene involved, is still not known.[27][28]

Alternative hypotheses

Several alternative hypotheses to natural selection as the driving force of evolution were proposed during the 1920s and 1930s. Random mutation, migration or genetic drift were also seen as major forces of evolution.[29]

P. A. Riley proposed an additional selective factor, where heavy metal chelation by melanin would supposedly protect peppered moths against the toxic effects of heavy metals associated with industrialisation. This selective advantage would supplement the major selective mechanism of differential bird predation.[30]

Phenotypic induction

In 1920, John William Heslop-Harrison rejected Tutt's differential bird predation hypothesis, on the basis that he did not believe that birds ate moths. Instead he proposed that pollutants could cause changes to the soma and germ plasm of the organism.[31] In 1925, K. Hasebroek made an early attempt to prove this hypothesis, exposing pupae to pollutant gases, namely hydrogen sulfide (H2S), ammonia (NH3) and "pyredin". He used eight species in his studies, four of which were species of butterfly that did not exhibit melanism.[32]

In 1926 and 1928, Heslop-Harrison suggested that the increase of melanic moths in industrialised regions was due to "mutation pressure", not to selection by predators which he regarded as negligible. Salts of lead and manganese were present in the airborne pollutant particles, and he suggested that these caused the mutation of genes for melanin production but of no others. He used Selenia bilunaria and Tephrosia bistortata as material. The larvae were fed with leaves that had incorporated these salts: melanics subsequently appeared.[33][34] A similar experiment in 1932 by McKenney Hughes failed to replicate these results; the statistician and geneticist Ronald Fisher showed that Heslop-Harrison's controls were inadequate, and that Hughes's findings made the 6% mutation rate required by Heslop-Harrison "improbable".[35]

Kettlewell's experiment

The first important experiments on the peppered moth were carried out by Bernard Kettlewell at Oxford University, under the supervision of E. B. Ford, who helped him gain a grant from the Nuffield Foundation to perform the experiments. In 1953, Kettlewell started a preliminary experiment in which moths were released into a large (18m × 6m) aviary, where they were fed on by great tits (Parus major). His main experiment, at Christopher Cadbury Wetland Reserve in Birmingham, England, involved marking, releasing and recapturing marked moths. He found that in this polluted woodland typica moths were preferentially preyed upon. He thus showed that the melanic phenotype was important to the survival of peppered moths in such a habitat. Kettlewell repeated the experiment in 1955 in unpolluted woodlands in Dorset, and again in the polluted woods in Birmingham.[3][23] In 1956 he repeated the experiments and found similar results; in Birmingham, birds ate most of the white moths (75%), whereas in Dorset, most of the dark moths (86%) were eaten.[2][36]

Criticisms

Theodore David Sargent[a] performed experiments between 1965 and 1969, from which he concluded that it was not possible to reproduce Kettlewell's results, and said that birds showed no preference for moths on either black or white tree trunks.[38][39] He suggested that Kettlewell had trained the birds to pick moths on tree trunks to obtain the desired results.[4][40]

Two chapters in Michael Majerus's 1998 book Melanism: Evolution in Action critiqued the research in Kettlewell's The Evolution of Melanism, discussed studies which raised questions about Kettlewell's original experimental methods, and called for further research.[23] Reviewing the book, Jerry Coyne noted these points, and concluded that "for the time being we must discard Biston as a well-understood example of natural selection in action, although it is clearly a case of evolution. There are many studies more appropriate for use in the classroom."[41]

Judith Hooper's book Of Moths and Men (2002) severely criticised Kettlewell's experiment.[42] Hooper argued that Kettlewell's field notes could not be found and suggested that his experiment was fraudulent, on the basis of Sargent's criticisms alleging that the photographs of the moths were taken of dead moths placed on a log. She said that E. B. Ford was a "Darwinian zealot",[43] and claimed that he exploited the scientifically naive Kettlewell to obtain the desired experimental results.[44] The book's reception led to demands that the peppered moth evolution story be deleted from textbooks.[45][46] Scientists have examined the allegations made by Hooper, and found them to be without merit.[22][47][48]

 
Creationists have disputed the occurrence or significance of the melanic carbonaria morph's increase in frequency.

Phillip E. Johnson, a co-founder of the creationist intelligent design movement, said that the moths "do not sit on tree trunks", that "moths had to be glued to the trunks" for pictures, and that the experiments were "fraudulent" and a "scam."[49] The intelligent design advocate Jonathan Wells wrote an essay on the subject, a shortened version of which appeared in the 24 May 1999 issue of The Scientist, claiming that "The fact that peppered moths do not normally rest on tree trunks invalidates Kettlewell's experiments".[50] Wells further wrote in his 2000 book Icons of Evolution that "What the textbooks don't explain, however, is that biologists have known since the 1980s that the classical story has some serious flaws. The most serious is that peppered moths in the wild don't even rest on tree trunks. The textbook photographs, it turns out, have been staged."[51] However, peppered moths do rest on tree trunks on occasion, and Nick Matzke states that there is little difference between the 'staged' photos and 'unstaged' ones.[52]

Majerus's experiment

From 2001 to 2007, Majerus carried out experiments in Cambridge to resolve the various criticisms of Kettlewell's experiment. During his experiment, he noted the natural resting positions of peppered moths. Of the 135 moths examined, over half were on tree branches, mostly on the lower half of the branch; 37% were on tree trunks, mostly on the north side; and only 12.6% were resting on or under twigs. Following correspondence with Hooper, he added an experiment to find if bats, not birds, could be the main predators. He observed a number of species of bird actually preying on the moths, and found that differential bird predation was a major factor responsible for the decline in carbonaria frequency compared to typica.[23] He described his results as a complete vindication of the natural selection theory of peppered moth evolution, and said "If the rise and fall of the peppered moth is one of the most visually impacting and easily understood examples of Darwinian evolution in action, it should be taught. It provides after all the proof of evolution."[53]

Majerus died before he could complete the writing up of his experiments, so the work was carried on by Cook, Grant, Saccheri and James Mallet, and published on 8 February 2012 as "Selective bird predation on the peppered moth: the last experiment of Michael Majerus."[54] The experiment became the largest ever in the study of industrial melanism, involving 4,864 individuals in a six-year investigation, and it confirmed that melanism in moths is a genuine example of natural selection involving camouflage and predation. Their concluding remark runs: "These data provide the most direct evidence yet to implicate camouflage and bird predation as the overriding explanation for the rise and fall of melanism in moths."[11]

Coyne said he was "delighted to agree with this conclusion [of Majerus's experiment], which answers my previous criticisms about the Biston story."[55]

See also

Notes

  1. ^ Sargent (1936–2018) was a biologist at the University of Massachusetts Amherst.[37]

References

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Further reading

External links

  • Bruce Grant has written several papers on melanism in the peppered moth which are listed on his home page.
  • Online lecture: presented by Laurence Cook.
  • Young, Matt. "Moonshine: Why the Peppered Moth remains an Icon of Evolution". Department of Physics, Colorado School of Mines. Retrieved 3 January 2009.
  • . This is the transcript of Michael Majerus' lecture delivered to the British Humanist Association on Darwin Day 2004.
  • . This is the transcript of Majerus' lecture given at the European Society for Evolutionary Biology meeting on 23 August 2007. The accompanying PowerPoint presentation is .
  • Majerus, Michael E. N. (2009). "Industrial Melanism in the Peppered Moth, Biston betularia: An Excellent Teaching Example of Darwinian Evolution in Action". Evolution: Education and Outreach. 2 (1): 63–74. doi:10.1007/s12052-008-0107-y. Accusations of data fudging and scientific fraud in the case are found to be vacuous.
  • On 19 June 2009, published on this evolutionary phenomenon and implored UK readers to visit the Moths Count website and record their observations of local moths, in an effort to help increase the available data for researchers.

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This article is about the peppered moth s significance in evolutionary biology For its evolutionary ancestry see Insect evolution The evolution of the peppered moth is an evolutionary instance of directional colour change in the moth population as a consequence of air pollution during the Industrial Revolution The frequency of dark coloured moths increased at that time an example of industrial melanism Later when pollution was reduced the light coloured form again predominated Industrial melanism in the peppered moth was an early test of Charles Darwin s natural selection in action and it remains a classic example in the teaching of evolution 1 2 3 In 1978 Sewall Wright described it as the clearest case in which a conspicuous evolutionary process has actually been observed 4 5 Biston betularia f typica the white bodied peppered moth Biston betularia f carbonaria the black bodied peppered moth The dark coloured or melanic form of the peppered moth var carbonaria was rare though a specimen had been collected by 1811 After field collection in 1848 from Manchester an industrial city in England the frequency of the variety was found to have increased drastically By the end of the 19th century it almost completely outnumbered the original light coloured type var typica with a record of 98 in 1895 6 The evolutionary importance of the moth was only speculated upon during Darwin s lifetime It was 14 years after Darwin s death in 1896 that J W Tutt presented it as a case of natural selection 7 Because of this the idea spread widely and more people came to believe in Darwin s theory Bernard Kettlewell was the first to investigate the evolutionary mechanism behind peppered moth adaptation between 1953 and 1956 He found that a light coloured body was an effective camouflage in a clean environment such as in rural Dorset while the dark colour was beneficial in a polluted environment like industrial Birmingham This selective survival was due to birds which easily caught dark moths on clean trees and white moths on trees darkened with soot The story supported by Kettlewell s experiment became the canonical example of Darwinian evolution and evidence for natural selection used in standard textbooks 8 9 10 However failure to replicate the experiment and Theodore David Sargent s criticism of Kettlewell s methods in the late 1960s led to general skepticism When Judith Hooper s Of Moths and Men was published in 2002 Kettlewell s story was more sternly attacked and accused of fraud The criticism became a major argument for creationists Michael Majerus was the principal defender His seven year experiment beginning in 2001 the most elaborate of its kind in population biology the results of which were published posthumously in 2012 vindicated Kettlewell s work in great detail This restored peppered moth evolution as the most direct evidence and one of the clearest and most easily understood examples of Darwinian evolution in action 11 Contents 1 Origin and evolution 1 1 Rise and fall of phenotype frequency 2 Genetics 3 Alternative hypotheses 3 1 Phenotypic induction 4 Kettlewell s experiment 4 1 Criticisms 5 Majerus s experiment 6 See also 7 Notes 8 References 9 Further reading 10 External linksOrigin and evolution Edit Typica and carbonaria morphs on the same tree The light coloured typica below the bark s scar is nearly invisible on this pollution free tree camouflaging it from predators Further information Industrial melanism Before the Industrial Revolution the black form of the peppered moth was rare The first black specimen of unknown origin was collected before 1811 and kept in the University of Oxford 12 13 14 The first live specimen was caught by R S Edleston in Manchester England in 1848 but he reported this only 16 years later in 1864 in The Entomologist 15 Edleston notes that by 1864 it was the more common type of moth in his garden in Manchester The light bodied moths were able to blend in with the light coloured lichens and tree bark and the less common black moths were more likely to be eaten by birds As a result of the common light coloured lichens and English trees therefore the light coloured moths were much more effective at hiding from predators and the frequency of the dark allele was very low at about 0 01 16 During the early decades of the Industrial Revolution in England the countryside between London and Manchester became blanketed with soot from the new coal burning factories Many of the light bodied lichens died from sulphur dioxide emissions and the trees became darkened This led to an increase in bird predation for light coloured moths as they no longer blended in as well in their polluted ecosystem indeed their bodies now dramatically contrasted with the colour of the bark Dark coloured moths on the other hand were camouflaged very well by the blackened trees 17 The population of dark coloured moth rapidly increased By the mid 19th century the number of dark coloured moths had risen noticeably and by 1895 the percentage of dark coloured moths in Manchester was reported at 98 a dramatic change of almost 100 from the original frequency 17 This effect of industrialization in body colour led to the coining of the term industrial melanism 3 The implication that industrial melanism could be evidence supporting Charles Darwin s theory of natural selection was noticed during his lifetime Albert Brydges Farn 1841 1921 a British entomologist wrote to Darwin on 18 November 1878 to discuss his observation of colour variations in the Annulet moth then Gnophos obscurata now Charissa obscurata He noted the existence of dark moths in peat in the New Forest brown moths on clay and red soil in Herefordshire and white moths on chalk cliffs in Lewes and suggested that this variation was an example of survival of the fittest He told Darwin that he had found dark moths on a chalk slope where the foliage had been blackened by smoke from lime kilns and he had also heard that white moths had become less common at Lewes after lime kilns had been in operation for a few years 18 Darwin does not seem to have responded to this information possibly because he thought natural selection would be a much slower process 19 A scientific explanation of moth coloration was only published in 1896 14 years after Darwin s death when J W Tutt explicitly linked peppered moth melanism to natural selection 16 Rise and fall of phenotype frequency Edit Melanism has appeared in both European and North American peppered moth populations Information about the rise in frequency is scarce Much more is known about the subsequent fall in phenotype frequency as it has been measured by lepidopterists using moth traps Steward compiled data for the first recordings of the peppered moth by locality and deduced that the carbonaria morph was the result of a single mutation that subsequently spread By 1895 it had reached a reported frequency of 98 in Manchester 20 From around 1962 to the present the phenotype frequency of carbonaria has steadily fallen in line with cleaner air around industrial cities Its decline has been measured more accurately than its rise through more rigorous scientific studies Notably Kettlewell conducted a national survey in 1956 Bruce Grant conducted a similar one in early 1996 21 and L M Cook in 2003 22 Similar results were found in North America Melanic forms have not been found in Japan It is believed that this is because peppered moths in Japan do not inhabit industrialised regions 22 Genetics EditTutt was the first to propose the differential bird predation hypothesis in 1896 as a mechanism of natural selection The melanic morphs were better camouflaged against the bark of trees without foliose lichen whereas the typica morphs were better camouflaged against trees with lichens As a result birds would find and eat those morphs that were not camouflaged with increased frequency 23 In 1924 J B S Haldane calculated using a simple general selection model the selective advantage necessary for the recorded natural evolution of peppered moths based on the assumption that in 1848 the frequency of dark coloured moths was 2 and by 1895 it was 95 The dark coloured or melanic form would have had to be 50 more fit than the typical light coloured form Even taking into consideration possible errors in the model this reasonably excluded the stochastic process of genetic drift because the changes were too fast 24 Haldane s statistical analysis of selection for the melanic variant in peppered moths became a well known part of his effort to demonstrate that mathematical models that combined natural selection with Mendelian genetics could explain evolution an effort that played a key role in the foundation of the discipline of population genetics and the beginnings of the modern synthesis of evolutionary theory with genetics 25 The peppered moth Biston betularia is also a model of parallel evolution in the incidence of melanism in the British form f carbonaria and the American form f swettaria as they are indistinguishable in appearance Genetic analysis indicates that both phenotypes are inherited as autosomal dominants Cross hybridizations indicate that the phenotypes are produced by alleles at a single locus 26 The gene for carbonaria in B betularia was thought to be in a region of chromosome 17 It was later concluded that the gene could not be in that region because none of the genes in the chromosome coded for either wing pattern or melanisation The region that was used to find it was the first intron of the orthologue of the cortex gene in Drosophila Through elimination of candidates within the region based on rarity a 21 925 base pair insert remained The insert labelled carb TE is a class II transposable element that has an approximately 9 kb non repetitive sequence tandemly repeated two and one third times There are 6 base pairs of inverted repeats and duplicated 4 base pairs at the target site not present in typica moths Carb TE has higher expression during the stage of rapid wing disc morphogenesis The mechanism of how the gene increases expression and whether it is the only gene involved is still not known 27 28 Alternative hypotheses EditSeveral alternative hypotheses to natural selection as the driving force of evolution were proposed during the 1920s and 1930s Random mutation migration or genetic drift were also seen as major forces of evolution 29 P A Riley proposed an additional selective factor where heavy metal chelation by melanin would supposedly protect peppered moths against the toxic effects of heavy metals associated with industrialisation This selective advantage would supplement the major selective mechanism of differential bird predation 30 Phenotypic induction Edit In 1920 John William Heslop Harrison rejected Tutt s differential bird predation hypothesis on the basis that he did not believe that birds ate moths Instead he proposed that pollutants could cause changes to the soma and germ plasm of the organism 31 In 1925 K Hasebroek made an early attempt to prove this hypothesis exposing pupae to pollutant gases namely hydrogen sulfide H2S ammonia NH3 and pyredin He used eight species in his studies four of which were species of butterfly that did not exhibit melanism 32 In 1926 and 1928 Heslop Harrison suggested that the increase of melanic moths in industrialised regions was due to mutation pressure not to selection by predators which he regarded as negligible Salts of lead and manganese were present in the airborne pollutant particles and he suggested that these caused the mutation of genes for melanin production but of no others He used Selenia bilunaria and Tephrosia bistortata as material The larvae were fed with leaves that had incorporated these salts melanics subsequently appeared 33 34 A similar experiment in 1932 by McKenney Hughes failed to replicate these results the statistician and geneticist Ronald Fisher showed that Heslop Harrison s controls were inadequate and that Hughes s findings made the 6 mutation rate required by Heslop Harrison improbable 35 Kettlewell s experiment EditMain article Kettlewell s experiment The first important experiments on the peppered moth were carried out by Bernard Kettlewell at Oxford University under the supervision of E B Ford who helped him gain a grant from the Nuffield Foundation to perform the experiments In 1953 Kettlewell started a preliminary experiment in which moths were released into a large 18m 6m aviary where they were fed on by great tits Parus major His main experiment at Christopher Cadbury Wetland Reserve in Birmingham England involved marking releasing and recapturing marked moths He found that in this polluted woodland typica moths were preferentially preyed upon He thus showed that the melanic phenotype was important to the survival of peppered moths in such a habitat Kettlewell repeated the experiment in 1955 in unpolluted woodlands in Dorset and again in the polluted woods in Birmingham 3 23 In 1956 he repeated the experiments and found similar results in Birmingham birds ate most of the white moths 75 whereas in Dorset most of the dark moths 86 were eaten 2 36 Criticisms Edit Further information Kettlewell s experiment Criticisms Theodore David Sargent a performed experiments between 1965 and 1969 from which he concluded that it was not possible to reproduce Kettlewell s results and said that birds showed no preference for moths on either black or white tree trunks 38 39 He suggested that Kettlewell had trained the birds to pick moths on tree trunks to obtain the desired results 4 40 Two chapters in Michael Majerus s 1998 book Melanism Evolution in Action critiqued the research in Kettlewell s The Evolution of Melanism discussed studies which raised questions about Kettlewell s original experimental methods and called for further research 23 Reviewing the book Jerry Coyne noted these points and concluded that for the time being we must discard Biston as a well understood example of natural selection in action although it is clearly a case of evolution There are many studies more appropriate for use in the classroom 41 Judith Hooper s book Of Moths and Men 2002 severely criticised Kettlewell s experiment 42 Hooper argued that Kettlewell s field notes could not be found and suggested that his experiment was fraudulent on the basis of Sargent s criticisms alleging that the photographs of the moths were taken of dead moths placed on a log She said that E B Ford was a Darwinian zealot 43 and claimed that he exploited the scientifically naive Kettlewell to obtain the desired experimental results 44 The book s reception led to demands that the peppered moth evolution story be deleted from textbooks 45 46 Scientists have examined the allegations made by Hooper and found them to be without merit 22 47 48 Creationists have disputed the occurrence or significance of the melanic carbonaria morph s increase in frequency Phillip E Johnson a co founder of the creationist intelligent design movement said that the moths do not sit on tree trunks that moths had to be glued to the trunks for pictures and that the experiments were fraudulent and a scam 49 The intelligent design advocate Jonathan Wells wrote an essay on the subject a shortened version of which appeared in the 24 May 1999 issue of The Scientist claiming that The fact that peppered moths do not normally rest on tree trunks invalidates Kettlewell s experiments 50 Wells further wrote in his 2000 book Icons of Evolution that What the textbooks don t explain however is that biologists have known since the 1980s that the classical story has some serious flaws The most serious is that peppered moths in the wild don t even rest on tree trunks The textbook photographs it turns out have been staged 51 However peppered moths do rest on tree trunks on occasion and Nick Matzke states that there is little difference between the staged photos and unstaged ones 52 Majerus s experiment EditFurther information Michael E N Majerus From 2001 to 2007 Majerus carried out experiments in Cambridge to resolve the various criticisms of Kettlewell s experiment During his experiment he noted the natural resting positions of peppered moths Of the 135 moths examined over half were on tree branches mostly on the lower half of the branch 37 were on tree trunks mostly on the north side and only 12 6 were resting on or under twigs Following correspondence with Hooper he added an experiment to find if bats not birds could be the main predators He observed a number of species of bird actually preying on the moths and found that differential bird predation was a major factor responsible for the decline in carbonaria frequency compared to typica 23 He described his results as a complete vindication of the natural selection theory of peppered moth evolution and said If the rise and fall of the peppered moth is one of the most visually impacting and easily understood examples of Darwinian evolution in action it should be taught It provides after all the proof of evolution 53 Majerus died before he could complete the writing up of his experiments so the work was carried on by Cook Grant Saccheri and James Mallet and published on 8 February 2012 as Selective bird predation on the peppered moth the last experiment of Michael Majerus 54 The experiment became the largest ever in the study of industrial melanism involving 4 864 individuals in a six year investigation and it confirmed that melanism in moths is a genuine example of natural selection involving camouflage and predation Their concluding remark runs These data provide the most direct evidence yet to implicate camouflage and bird predation as the overriding explanation for the rise and fall of melanism in moths 11 Coyne said he was delighted to agree with this conclusion of Majerus s experiment which answers my previous criticisms about the Biston story 55 See also EditPolymorphism Scottish red deerNotes Edit Sargent 1936 2018 was a biologist at the University of Massachusetts Amherst 37 References Edit Cook L M Saccheri I J March 2013 The peppered moth and industrial melanism evolution of a natural selection case study Heredity 110 3 207 212 doi 10 1038 hdy 2012 92 PMC 3668657 PMID 23211788 a b Rudge David W 2005 The Beauty of Kettlewell s Classic Experimental Demonstration of Natural Selection BioScience 55 4 369 375 doi 10 1641 0006 3568 2005 055 0369 TBOKCE 2 0 CO 2 a b c Majerus Michael E N 2008 Industrial Melanism in the Peppered Moth Biston betularia An Excellent Teaching Example of Darwinian Evolution in Action PDF Evolution Education and Outreach 2 1 63 74 doi 10 1007 s12052 008 0107 y S2CID 25407417 a b Rice Stanley A 2007 Encyclopedia of Evolution New York Facts on File p 308 ISBN 978 1 4381 1005 9 Majerus Michael E N 1998 Melanism Evolution in Action Oxford University Press ISBN 978 0 19 854983 3 Clarke C A Mani G S Wynne G 1985 Evolution in reverse clean air and the peppered moth Biological Journal of the Linnean Society 26 2 189 199 doi 10 1111 j 1095 8312 1985 tb01555 x Majerus Michael E N 2008 Industrial Melanism in the Peppered Moth Biston betularia An Excellent Teaching Example of Darwinian Evolution in Action Evolution Education and Outreach 2 1 63 74 doi 10 1007 s12052 008 0107 y Bender Eric 21 March 2022 Urban evolution How species adapt to survive in cities Knowable Magazine Annual Reviews doi 10 1146 knowable 031822 1 Diamond Sarah E Martin Ryan A 2 November 2021 Evolution in Cities Annual Review of Ecology Evolution and Systematics 52 1 519 540 doi 10 1146 annurev ecolsys 012021 021402 S2CID 239646134 Miller Ken 1999 The Peppered Moth An Update www millerandlevine com Retrieved 31 March 2022 a b Cook L M Grant B S Saccheri I J Mallet James 2012 Selective bird predation on the peppered moth the last experiment of Michael Majerus Biology Letters 8 4 609 612 doi 10 1098 rsbl 2011 1136 PMC 3391436 PMID 22319093 Berry R J 1990 Industrial melanism and peppered moths Biston betularia L Biological Journal of the Linnean Society 39 4 301 322 doi 10 1111 j 1095 8312 1990 tb00518 x Saccheri I J Rousset F Watts P C Brakefield P M Cook L M 2008 Selection and gene flow on a diminishing cline of melanic peppered moths PNAS 105 42 16212 16217 Bibcode 2008PNAS 10516212S doi 10 1073 pnas 0803785105 PMC 2571026 PMID 18854412 Neal Dick 2004 Introduction to Population Biology Reprint ed Cambridge Cambridge University Press p 171 ISBN 9780521532235 Edleston R S 1864 No title Entomologist 2 150 a b Hart Adam G Stafford Richard Smith Angela L Goodenough Anne E 2010 Evidence for contemporary evolution during Darwin s lifetime PDF Current Biology 20 3 R95 doi 10 1016 j cub 2009 12 010 PMID 20144776 S2CID 31093691 a b Miller Ken 1999 The Peppered Moth An Update Farn A B 18 November 1878 Farn A B to Darwin C R Darwin Correspondence Project Letter 11747 The Darwin Papers Manuscripts Room Cambridge University Library West Road Cambridge England DAR 164 26 How an extraordinary letter to Darwin spotted industrial melanism in moths Science Focus 27 February 2018 Retrieved 16 June 2018 Steward R C 1977 Industrial and non industrial melanism in the peppered moth Biston betularia L Ecological Entomology 2 3 231 243 doi 10 1111 j 1365 2311 1977 tb00886 x S2CID 85624115 Grant Bruce Distribution of melanism in Britain Talk Origins Retrieved 2 June 2022 a b c Cook L M 2003 The rise and fall of the Carbonaria form of the peppered moth The Quarterly Review of Biology 78 4 399 417 doi 10 1086 378925 PMID 14737825 S2CID 26831926 a b c d Majerus Michael E N August 2007 The Peppered Moth The Proof of Darwinian Evolution PDF Archived from the original PDF on 15 June 2011 Retrieved 9 September 2007 Haldane J B S 1924 A Mathematical Theory of Natural and Artificial Selection Bowler Peter J 2003 Evolution The history of an idea 3rd edition University of California Press pp 331 332 ISBN 0 520 23693 9 Grant B S 2004 Allelic melanism in American and British peppered moths Journal of Heredity 95 2 97 102 doi 10 1093 jhered esh022 PMID 15073224 van t Hof Arjen E Edmonds Nicola Dalikova Martina Marec Frantisek Saccheri Ilik J 20 May 2011 Industrial Melanism in British Peppered Moths Has a Singular and Recent Mutational Origin Science 332 6032 958 960 Bibcode 2011Sci 332 958V doi 10 1126 science 1203043 PMID 21493823 S2CID 24400858 via JSTOR van t Hof Arjen E Campagne Pascal Rigden Daniel J Yung Carl J Lingley Jessica Quail Michael A Hall Neil Darby Alistair Saccheri Ilik J 2 June 2016 The industrial melanism mutation in British peppered moths is a transposable element Nature 534 7605 102 117 Bibcode 2016Natur 534 102H doi 10 1038 nature17951 PMID 27251284 S2CID 3989607 Dobzhansky T G 1937 Genetics and the Evolutionary Process Columbia University Press ISBN 0 231 08306 8 Riley P A 2013 A proposed selective mechanism based on metal chelation in industrial melanic moths PDF Biological Journal of the Linnean Society 109 2 298 301 doi 10 1111 bij 12062 Heslop Harrison J W 1920 Genetical studies in the moths of the geometrid genus Oporabia Oporinia with a special consideration of melanism in the lepidoptera Journal of Genetics 9 3 195 280 doi 10 1007 BF02983273 S2CID 38996034 Hasebroek K 1925 Die prinzipielle Loesung des Problems des Grossstadt und Industriemelanismus der Schmetterlinge The principal solution of problems of city and industrial melanism of butterflies Internationale entomologische Zeitschrift in German 19 Heslop Harrison J W Garrett F C 1926 The induction of melanism in the lepidoptera and its subsequent inheritance Proceedings of the Royal Society of London Series B 99 696 241 263 rspb 1926 0012 doi 10 1098 rspb 1926 0012 ISSN 0950 1193 S2CID 84987348 Heslop Harrison J W 1928 A further induction of melanism in the lepidopterous insect Selenia bilunaria Esp and its inheritance Proceedings of the Royal Society of London Series B 102 718 338 347 doi 10 1098 rspb 1928 0009 Fisher R A 1933 On the Evidence Against the Chemical Induction of Melanism in Lepidoptera Proceedings of the Royal Society B 112 778 407 416 Bibcode 1933RSPSB 112 407F doi 10 1098 rspb 1933 0018 Kettlewell H B D 1958 A survey of the frequencies of Biston betularia L Lep and its melanic forms in Great Britain Heredity 12 1 51 72 doi 10 1038 hdy 1958 4 Obituary Theodore Sargent Professor Emeritus of Biology University of Massachusetts Amherst 4 September 2018 Retrieved 31 May 2022 Sargent T D 1968 Cryptic moths effects on background selections of painting the circumocular scales Science 159 3810 100 101 Bibcode 1968Sci 159 100S doi 10 1126 science 159 3810 100 PMID 5634373 S2CID 32124765 Sargent T D 1969 Background Selections of the Pale and Melanic Forms of the Cryptic Moth Phigalia titea Cramer Nature 222 5193 585 586 Bibcode 1969Natur 222 585S doi 10 1038 222585b0 S2CID 4202131 Sargent T D Millar C D Lambert D M 1988 Chapter 9 The classical explanation of industrial melanism Assessing the evidence In Hecht Max K Wallace Bruce eds Evolutionary Biology Vol 23 Plenum Press ISBN 0306429772 Coyne Jerry A 1998 Not Black and White Review of Melanism Evolution in Action by Michael E N Majerus Nature 396 6706 35 36 doi 10 1038 23856 Kenney Michael 22 October 2002 Of Dark Moths Men and Evolution Chicago Tribune Retrieved 10 December 2014 Of Moths and Men W W Norton amp Company Archived from the original on 10 December 2014 Retrieved 10 December 2014 Smith Peter D 11 May 2002 Of Moths and Men Intrigue Tragedy amp the Peppered Moth The Guardian Retrieved 10 December 2014 Of moths and men The Independent 4 September 2003 Archived from the original on 25 May 2022 Retrieved 10 December 2014 Dover Gabby 2003 Mothbusters EMBO Reports 4 3 235 doi 10 1038 sj embor embor778 PMC 1315906 Grant B S 2002 Sour grapes of wrath Science 297 5583 940 941 doi 10 1126 science 1073593 S2CID 161367302 Majerus Michael E N 2005 The peppered moth decline of a Darwinian disciple In Fellowes Mark Holloway Graham Rolf Jens eds Insect Evolutionary Ecology Wallingford Oxon CABI Publishing pp 375 377 ISBN 978 1 84593 140 7 Frack Donald 16 April 1999 Evolution April 1999 Peppered Moths and Creationists Archived from the original on 26 August 2007 Retrieved 26 August 2007 Wells J 24 May 1999 Second Thoughts about Peppered Moths This classical story of evolution by natural selection needs revising The Scientist 13 11 13 Wells J 2000 Icons of Evolution Science or Myth Why Much of What We Teach About Evolution is Wrong Regnery Press Washington D C p 138 book available from Iconsofevolution com Matzke Nicholas Icon of Obfuscation Chapter 7 Peppered Moths Connor Steve 25 August 2007 Moth study backs classic test case for Darwin s theory The Independent Archived from the original on 7 October 2008 Retrieved 9 September 2007 Matzke Nick 8 February 2012 Selective bird predation on the peppered moth the last experiment of Michael Majerus The Panda s Thumb Retrieved 7 March 2012 Coyne Jerry 12 February 2012 The peppered moth story is solid Why Evolution Is True Retrieved 7 March 2012 Further reading EditMajerus Michael E N 1998 Melanism Evolution in Action Oxford Oxford University Press ISBN 978 0 198 54983 3 External links Edit Wikimedia Commons has media related to Biston betularia Bruce Grant has written several papers on melanism in the peppered moth which are listed on his home page Online lecture The rise and fall of the melanic Peppered Moth presented by Laurence Cook Young Matt Moonshine Why the Peppered Moth remains an Icon of Evolution Department of Physics Colorado School of Mines Retrieved 3 January 2009 The Peppered Moth Decline of a Darwinian Disciple This is the transcript of Michael Majerus lecture delivered to the British Humanist Association on Darwin Day 2004 The Peppered Moth The Proof of Darwinian Evolution This is the transcript of Majerus lecture given at the European Society for Evolutionary Biology meeting on 23 August 2007 The accompanying PowerPoint presentation is also available Majerus Michael E N 2009 Industrial Melanism in the Peppered Moth Biston betularia An Excellent Teaching Example of Darwinian Evolution in Action Evolution Education and Outreach 2 1 63 74 doi 10 1007 s12052 008 0107 y Accusations of data fudging and scientific fraud in the case are found to be vacuous On 19 June 2009 Telegraph co uk published an article on this evolutionary phenomenon and implored UK readers to visit the Moths Count website and record their observations of local moths in an effort to help increase the available data for researchers Retrieved from https en wikipedia org w index php title Peppered moth evolution amp oldid 1168181711, wikipedia, wiki, book, books, library,

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