Peppered moth evolution: Difference between revisions
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{{about|the peppered moth's significance in [[evolutionary biology]]|its evolutionary ancestry|Insect evolution}} |
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[[Image:Biston.betularia.7200.jpg|thumb|260px|''Biston betularia f. typica'', the white-bodied [[peppered moth]].]] |
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[[Image:Biston.betularia.f.carbonaria.7209.jpg|thumb|260px|''Biston betularia f. carbonaria'', the black-bodied peppered moth.]] |
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The '''evolution of the [[peppered moth]]''' over the last two hundred years has been studied in detail. Originally, the vast majority of peppered moths had light colouration, which effectively [[camouflage]]d them against the light-coloured trees and lichens which they rested upon. However, because of widespread pollution during the [[Industrial Revolution]] in England, many of the lichens died out, and the trees that peppered moths rested on became blackened by [[soot]], causing most of the light-coloured moths, or '''''typica''''', to die off from predation. At the same time, the dark-coloured, or ''melanic'', moths, '''''carbonaria''''', flourished because of their ability to hide on the darkened trees.<ref name="miller">Miller, Ken (1999). ''[http://www.millerandlevine.com/km/evol/Moths/moths.html The Peppered Moth: An Update]''</ref> |
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Since then, with improved environmental standards, light-coloured peppered moths have again become common, but the dramatic change in the peppered moth's population has remained a subject of much interest and study, and has led to the coining of the term '''''industrial [[melanism]]''''' to refer to the genetic darkening of species in response to pollutants. As a result of the relatively simple and easy-to-understand circumstances of the adaptation, the peppered moth has become a common example used in explaining or demonstrating [[natural selection]].<ref name="globalchange">[http://web.archive.org/web/19960101-re_/http://www.globalchange.umich.edu/globalchange1/current/labs/peppered_moth/pepperedmoth.html A modelling exercise for students using the peppered moth as its example]</ref> |
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== Genetics == |
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{{main|Peppered moth|genetics}} |
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[[Evolution]] is defined as "a change in the frequency of an [[allele]] within a [[gene pool]]",<ref name="Dobzhansky">Dobzhansky, T.G. (1937). ''Genetics and the Evolutionary Process''. Columbia University Press. ISBN 0-231-08306-8.</ref> an occurrence that causes a [[population genetics|population]]'s genetically inherited [[trait (biology)|traits]] to change over successive generations. Evolution in the wild is chiefly caused by two mechanisms: [[natural selection]], the process by which individual organisms with beneficial traits are more likely to survive and reproduce, and [[genetic drift]], the statistical drift over time of [[allele frequency|allele frequencies]] in a population from random sampling effects in the formation of successive generations. |
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[[J.W. Tutt]] first proposed 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. |
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In 1924, [[J.B.S. Haldane]] calculated, using a simple [[general selection model]], the selective advantage necessary for the recorded 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 one and a half times as fit as the typical, light-coloured form. Even taking into consideration the errors in the model, this reasonably excluded the [[stochastic]] process of genetic drift, because the changes were too fast.<ref name="haldane">[[J. B. S. Haldane|Haldane, J.B.S.]] (1924). ''[[A Mathematical Theory of Natural and Artificial Selection]]''.</ref> |
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In peppered moths, the allele for dark-bodied moths is [[dominant allele|dominant]], while the allele for light-bodied moths is [[recessive gene|recessive]], meaning that the ''typica'' moths have a [[phenotype]] (visible or detectable characteristic) that is only seen in a [[homozygote|homozygous]] [[genotype]] (an organism that has two copies of the same allele), and never in a [[heterozygote|heterozygous]] one. This helps explain how dramatically quickly the population changed when being [[selection|selected]] for dark colouration. |
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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 dominant]]s. Cross hybridizations indicate the phenotypes are produced by isoalleles at a single locus.<ref>{{cite journal | author = Grant BS.| title = Allelic melanism in American and British peppered moths..| journal = Journal of Heredity| volume = 95 | issue =2| pages = 97–102 | year =2004 | pmid = 15073224 | doi = 10.1093/jhered/esh022}}</ref> |
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== Environmental changes == |
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[[File:Lichte en zwarte versie berkenspanner.jpg|thumb|360px|''Typica'' and ''carbonaria'' morphs resting on the same tree. The light-colored ''typica'' (below the bark's scar) is nearly invisible on this pollution-free tree, [[camouflage|camouflaging]] it from predators.]] |
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Before the [[Industrial Revolution]], the peppered moth was mostly found in a light grey form with little black speckled spots. The light-bodied moths were able to blend in with the light-coloured lichens and tree bark, and the less common black moth was 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 about 0.01%. |
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During the early decades of the Industrial Revolution in England, the countryside between London and Manchester was blanketed with [[soot]] from the new coal-burning factories. Many of the light-bodied lichens died from [[sulfur dioxide|sulphur dioxide]] emissions, and the trees became covered with soot. This led to an increase in bird predation for light-coloured moths, as they no longer blended in as well in their [[air pollution|polluted]] [[ecosystem]]: indeed, their bodies now dramatically contrasted with the colour of the bark. Dark-coloured moths, on the other hand, were [[camouflage]]d very well by the blackened trees.<ref name="miller"/> |
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Although a majority of light-coloured moths initially continued to be produced, most of them didn't survive, while the dark-coloured moths flourished. As a result, over the course of many generations of moths, the allele frequency gradually shifted towards the dominant allele, as more and more dark-bodied moths survived to reproduce. By the mid-19th century, the number of dark-coloured moths had risen noticeably, and by 1895, the percentage of dark-coloured moths in the Manchester peppered moth population was reported at 98%, a dramatic change (by almost 1000%) from the original frequency.<ref name="miller"/> This evolved darkening in colour as a result of industrialization has come to be known as ''industrial [[melanism]]'' as a result. |
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While evidence of increasing frequency of melanic forms in the Lepidoptera was available during Darwin’s lifetime – the first observations were made in 1848<ref name="Edleston">{{cite journal | last1 = Edleston | first1 = R.S. | year = 1864 | title = Amphydaris betularia | url = | journal = Entomologist | volume = 2 | issue = | page = 150 }}</ref> – current understanding is that it was not until 1896, 14 years after Darwin’s death, that Tutt explicitly linked melanism with natural selection. However, a recent article<ref name="Hart">{{cite journal | last1 = Hart | first1 = A.G. | last2 = Stafford | first2 = R. | last3 = Smith | first3 = A.L. | last4 = Goodenough | first4 = A.E. | year = 2010 | title = Evidence for contemporary evolution during Darwin's lifetime | url = | journal = Current Biology | volume = 20 | issue = 3| page = R95 }}</ref> reports that melanism in the Lepidoptera had been linked to natural selection prior to Tutt. Albert Brydges Farn (1841–1921), a British entomologist, wrote to Darwin on the 18th November 1878 to discuss his observation of colour variations in the Annulet moth (then ''Gnophos obscurata'', now ''Charissa obscurata''). In his letter, Farn<ref name="Farn">Farn, A.B. (1878). Farn, A.B. to Darwin C.R. 18 Nov 1878, Darwin Correspondence Project Letter 11747, held as DAR 164:26 as part of The Darwin Papers, Manuscripts Room, Cambridge University Library, West Road, Cambridge, England. http://www.darwinproject.ac.uk/entry-11747</ref> mentions the existence of different colour morphs, describing how each is matched to the habitats in which they are found (dark morphs on peat, white morphs on chalk cliffs) and refers explicitly to this variability as pointing to ‘survival of the fittest’. |
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In modern times, because of cleaner air standards in Europe and North America, the dark-bodied moth is becoming less frequent, again demonstrating the [[adaptation|adaptive]] shifts in the peppered moth population.<ref name="globalchange"/> |
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== Rise and fall of phenotype frequency == |
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[[Melanism]] has appeared in the 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 [[lepidopterist]]s using [[moth trapping|moth traps]]. |
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Though a black peppered moth was found in 1811, this can be seen as an aberrant morph caused by a recurrent mutation that was probably selected out of the population. The first ''carbonaria'' to be found was caught in [[Manchester]], [[England]] in 1848, but was only reported 16 years later in 1864 by Edleston. Edleston notes that by 2012 it was the more common morph in his garden in Manchester. 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% PEANUT BUTTER!!!!! |
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in Manchester.<ref name="steward">{{cite journal | last1 = Steward | first1 = R.C. | year = 1977 | title = Industrial and non-industrial melanism in the peppered moth ''Biston betularia'' (L.) | url = | journal = Ecological Entomology | volume = 2 | issue = 3| pages = 231–243 | doi = 10.1111/j.1365-2311.1977.tb00886.x }}</ref> |
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From around 1962 to the present, the phenotype frequency of ''carbonaria'' has steadily fallen. Its decline has been measured more accurately than its rise, because of more rigorous scientific studies being conducted. Notably, Bernard Kettlewell conducted a national survey in 1956, Bruce Grant conducted a similar one in early 1996.<ref>http://www.talkorigins.org/faqs/wells/images/grantfile.jpg</ref> |
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<!-- (figure 3, until permission can be granted (pun not intended), it will not be uploaded onto Wiki server) -->, and L.M. Cook in 2003.<ref>{{cite journal | author = Cook LM.| title = The rise and fall of the Carbonaria form of the peppered moth| journal = The Quarterly Review of Biology | issue =4| pages = 399–417 | year =2003 | pmid = 14737825 | volume = 78 | doi = 10.1086/378925}}</ref> |
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Similar results were found in 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. |
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==Predation experiments== |
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[[Image:Parusmajorde.jpg|thumb|left|200px|The [[Great Tit]], an insectivorous bird.]] |
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In 1896 [[J. W. Tutt]] hypothesised that the increased proportion of ''carbonaria'' resulted from differential bird [[predation]] giving an advantage to the melanistic phenotype in polluted regions, but not in unpolluted regions where the light coloured ''typica'' moths had the advantage.<ref name=swedentalk>{{cite web |url=http://www.gen.cam.ac.uk/research/personal/majerus/SwedenPepperedmoth2007Ppt.pdf |format=PDF|title=The Peppered Moth: The Proof of Darwinian Evolution |accessdate=2007-09-09 |author=Michael E. N. Majerus |authorlink= |coauthors= |month=August | year=2007 |
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|work= |publisher= |pages= |language= |archiveurl = http://www.gen.cam.ac.uk/research/personal/majerus/Swedentalk220807.pdf <!-- Bot retrieved archive --> |archivedate = 2008-02-21 |quote=}}</ref> Various experiments have been performed on predation of the peppered moth and each has supported this hypothesis. |
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The most famous experiments on the peppered moth were carried out by [[Bernard Kettlewell]] under the supervision of [[E. B. Ford]], who helped him gain a grant from the [[Nuffield Foundation]] to perform the experiments. In one of [[Kettlewell's experiment]]s, moths were released into a large (18 m by 6 m) aviary, where they were fed on by [[Great Tit]]s (''Parus major''). In 1953, Kettlewell experimented at [[Cadbury Nature Reserve]] in [[Birmingham]], [[England]], marking, releasing and recapturing marked moths. He found that in this polluted woodland ''typica'' morphs were preferentially preyed. He thus showed that the melanistic phenotype was important to the survival of peppered moths in such a habitat. Kettlewell repeated the experiment in 1955 at unpolluted woodland in [[Dorset]] and again in the polluted woods in Birmingham. He was accompanied by [[Nico Tinbergen]], and they made a film together. Further studies by others found similar results, culminating in 1996 when reporting work on both sides of the Atlantic found a correlation between changes in melanic frequencies and pollution levels.<ref name=swedentalk/><ref name="young">Young, M. (2003). ''[http://www.talkreason.org/articles/moonshine.cfm Moonshine: Why the Peppered Moth Remains an Icon of Evolution].''</ref> |
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An experiment in [[field biology]] will always suffer from some level of artificiality, but that has to be balanced against practicality, costs and in this case the history of field biology; the most important aspect is that an experiment generates useful statistics. The only previous experiments of this type were [[Ronald Fisher|R.A. Fisher]] and [[E.B. Ford]]'s (1947) with the [[scarlet tiger moth]]. |
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[[Michael Majerus]] in his 1998 book ''[[Melanism: Evolution in Action]]'' discussed criticisms concerning Kettlewell's experimental methods.<ref name=swedentalk/> [[#Criticism and controversy|Criticism and controversy]] arose when the book was misrepresented in reviews, and the story was picked up by [[creationism|creationist]] campaigners. The journalist [[Judith Hooper]] suggested in her book ''[[Of Moths and Men]]'' (2002) that Kettlewell committed scientific [[fraud]]. Careful studies of Kettlewell's surviving papers by Rudge (2005) and Young (2004) have revealed that Hooper's allegation of fraud is unjustified, and "that Hooper does not provide one shred of evidence to support this serious allegation”.<ref name="young"/><ref name="rudge2005">{{cite journal | last1 = Rudge | first1 = D.W. | year = 2005 | title = Did Kettlewell Commit Fraud? Re-examining the Evidence | url = | journal = Public Understanding of Science | volume = 14 | issue = 3| pages = 249–268 | doi = 10.1177/0963662505052890 | pmid = 16240545 }}</ref> |
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In 2000 Majerus developed plans for experiments to establish where peppered moths rest through the day, and to examine if the various valid criticisms of Kettlewell’s experimental protocols could have altered the qualitative validity of his conclusions. In the following year he piloted a new field predation experiment designed to overcome criticisms that Kettlewell had used too few release sites, resulting in the density of moths being too high; moths had been released onto tree trunks rather than branches; moths released during the day might not have found the best places to hide; mixtures of wild-caught and lab-bred moths might have behaved differently; and the behaviour of translocated moths might have changed because of local adaptation. During the main experiment in [[Cambridge]] over the seven years 2001-2007 Majerus noted the natural resting positions of the moths, and 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 [[bat]] predation might have skewed the results – this found that bats preyed equally on both forms of the moth. He observed a number of species of bird preying on the moths, and the overall data led him to conclude that differential bird predation was a major factor responsible for the decline in ''carbonaria'' frequency compared to ''typica'' in Cambridge during the study period.<ref name=swedentalk/> He described his results as a complete vindication of the peppered moth story, 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."<ref name=indy>{{cite web |url=http://news.independent.co.uk/sci_tech/article2893896.ece |title=Moth study backs classic 'test case' for Darwin's theory - Independent Online Edition > Sci_Tech |accessdate=2007-09-09 |author=Steve Connor, Science Editor |date=25 August 2007 |format= |work= |publisher=The Independent |pages= |language= |archiveurl= |archivedate= |quote=}}</ref> |
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==Alternative hypotheses== |
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Several alternative hypotheses to explain industrial [[melanism]], particularly noted in the peppered moth, were proposed during the 1920s and 1930s. Some dissenters within the scientific community have criticized the peppered moth story, notably Sargent ''et al.'' (1998), but peppered moth researchers remain unconvinced.<ref name="sargent">Sargent, T.D., Millar, C.D., and Lambert, D.M. (1998). "The 'classical' explanation of industrial melanism: Assessing the evidence.", Hecht and Wallace (Eds.), ''Evolutionary Biology'' volume 23, chapter 9.</ref> |
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Several alternative selection mechanisms have been proposed. Note that a change in [[allele frequency]], be it caused by [[natural selection]], [[mutation]], [[Insect migration|migration]] or [[genetic drift]] by definition, is [[cellular differentiation|differential]].<ref name="Dobzhansky"/> However, the magnitude of the changes observed can only be accounted for by natural selection. It can be seen from [[population genetics]] that a non-differential change will not cause evolution. If the allele frequencies are denoted by the [[algebra]]ic terms ''p'' and ''q'', and (say) ''p'' = 0.6 and ''q'' = 0.4, then a non-differential reduction in population size from say 2000 to 100 individuals, will still produce the same values of (approximately) ''p'' = 0.6 and ''q'' = 0.4. |
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===Phenotypic induction=== |
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[[John William Heslop-Harrison]] (1920) rejected Tutt's differential bird predation hypothesis, on the basis that he did not believe that birds ate moths. Instead he advocated the idea that pollutants could cause changes to the soma and germ plasm of the organism. |
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The origin of this hypothesis probably has its roots in the 1890s, when it was proposed as a form of [[Lamarckism]]. It is important to note its historical context. |
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Hasebroek (1925) was the first who tried to prove this hypothesis, <!-- reference? --> he contended that air pollution altered lepidopteran physiology, thus producing an excess of black pigment. He exposed pupae of Lepidoptera to various doses of pollutant gases, namely [[hydrogen sulfide]] (H<sub>2</sub>S), [[ammonia]] (NH<sub>3</sub>) and "pyredin" (presumably his spelling of [[pyridine]]). He used eight species in his studies, four of which were species of butterfly that did not exhibit melanism. Ford (1964) contends that Hasebroek's illustrations showed that the abnormal forms that appeared were not melanics, and Hasebroek failed to study their genetics. |
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[[John William Heslop-Harrison|Heslop Harrison]] (Harrison and Garrett 1926; Harrison 1928) 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 and melanics subsequently appeared. |
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Similar experiments by Hughes McKenney (1932) and Thomasen and Lemche (1933) failed to replicate these results. However, the statistician and geneticist [[Ronald Fisher]], showed that Heslop Harrison's controls were inadequate.<ref name="fisher">{{cite journal | last1 = Fisher | first1 = R.A. | year = 1933 | title = On the Evidence Against the Chemical Induction of Melanism in Lepidoptera | url = | journal = Proceedings of the Royal Society of London, B | volume = 112 | issue = 778| pages = 407–416 | doi = 10.1098/rspb.1933.0018 }}</ref> This hypothesis, however, appeared to be falsified by breeding experiments. Further evidence, if it were needed, is likely to come from research into the biochemistry of melanism. |
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==Criticism== |
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[[File:Ahlenmoor 3(loz).jpg|thumb|260px|[[Creationism|Creationists]] have disputed the occurrence or significance of the melanic ''carbonaria'' morph increasing in [[allele frequency|frequency]].]] |
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The use of the peppered moth as an example of evolution has come under attack by [[creationism|creationists]], including advocates of [[intelligent design]], who allege that it is not reliable as [[evidence of evolution]]. Creationists have argued that the "peppered moth story" showed only [[microevolution]], rather than [[speciation]] or other changes at the larger [[macroevolution]]ary scale.<ref>{{cite web |url=http://www.answersingenesis.org/articles/am/v3/n2/much-ado-about-moths |title=Much Ado About Moths |publisher=Answers in Genesis |author=Dr. Tommy Mitchell |date=2005-05-02}} (also [[Watchtower Bible and Tract Society]], 1985)</ref> Biologists agree that this example shows natural selection causing evolution within a species, demonstrating rapid and obvious adaptiveness with such change,<ref name=Obfuscation>{{cite web |url=http://www.talkorigins.org/faqs/wells/iconob.html#moths |title=Icon of Obfuscation |format= |work= |accessdate=2008-05-22}}</ref><ref>The Cambridge Dictionary of Human Biology and Evolution, p. 283, entry "Kettlewell, Henry B. D." Mai, Larry, Young Owl, Marcus & M Patricia Kersting. (eds.) Cambridge University Press.</ref> and accept that it is not proof of the theory of evolution as a whole. Although creationists accept microevolution of varieties within a kind, they claim that macroevolution does not happen. To biologists there is no dividing line between the two, and in the [[modern evolutionary synthesis]] the same mechanisms are seen operating at various scales to cause both evolution within species and speciation at a macroevolution level or wider changes, the only difference being of time and scale.<ref>[http://www.straightdope.com/mailbag/mmicromacroev.html The Straight Dope: Do creationists accept microevolution but not macroevolution?]</ref><ref name=Macroevolution>{{cite web |url=http://www.talkorigins.org/faqs/macroevolution.html#barriers |title=Macroevolution: Its definition, Philosophy and History |format= |work= |accessdate=2008-05-22}}</ref> |
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Another common creationist criticism involves well-known pictures of moths resting on trunks that are used in many textbooks. These photos were prepared (dead moths pinned to branches), which has been conflated into the idea that all the studies were staged. Professional photography to illustrate textbooks uses dead insects because of the considerable difficulty in getting good images of small, relatively fast moving animals. The studies actually consisted of observational data rather than using such photographs. The photographs in [[Michael Majerus]]'s 1998 book ''[[Melanism: Evolution in Action]]'' are unstaged pictures of live moths in the wild, and the photographs of moths on tree-trunks, apart from some slight blurring, look no different than the "staged" photographs.<ref name=ioo>{{cite web |url=http://www.talkorigins.org/faqs/wells/iconob.html |title=Icon of Obfuscation |accessdate=2007-08-25 |author=Nick Matzke |authorlink=Nick Matzke |date=2002-2004 |
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|work=Jonathan Wells's book ''Icons of Evolution'' and why most of what it teaches about evolution is wrong |publisher=[[TalkOrigins Archive]] |pages= |language= |archiveurl= |archivedate= |quote=}}</ref> While an experiment did involve the gluing of dead moths to trees, this practice was just one of many different ways used to study different individual elements of the overall hypothesis. This particular experiment was not meant to exactly reproduce natural conditions but instead was used to assess how the numbers of moths available (their density) affected the foraging practices of birds.<ref name=FraudMisleading>{{cite web |url=http://www.millerandlevine.com/km/evol/Moths/grant-pratt-tribune.html |title=Charges of Fraud Misleading |accessdate=2008-02-18 |author=Bruce Grant |authorlink=Bruce Grant |date=December 13, 2000 |work= |publisher=Pratt Tribune (Kansas) }}</ref> |
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The methodology of [[Bernard Kettlewell]]'s classic study was questioned by the biologist [[Jerry Coyne]] in ''Nature'' in a review of Michael Majerus's 1998 book ''Melanism: Evolution in Action'' which includes a critique of [[Kettlewell's experiment]], matching a similar 1998 analysis by Sargent ''et al.''<ref name="sargent"/> Coyne stated that the most serious problem found by Majerus was that only two peppered moths had been found on tree trunks. He also wrote that the white moths THEY EAT CHILDREN had increased in numbers before the lichen had returned and that Kettlewell's findings of moths choosing matching backgrounds had not been replicated in later experiments. Coyne compared his reaction to "the dismay attending my discovery, at the age of 6, that it was my father and not Santa who brought the presents on Christmas Eve". He 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" and that further studies of the animal's habits were needed.<ref name="coyne"> |
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{{cite journal |
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| last = Coyne |
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| first = Jerry A. |
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| coauthors = |
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| title = Not Black and White. Review of ''Melanism: Evolution in Action'' by Michael E.N. Majerus |
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| journal = Nature |
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| volume = 396 |
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| issue = 6706| pages = 35–36 |
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| date = 1998-11-05 |
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| publisher = |
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| url = http://pondside.uchicago.edu/ceb/Majerus_review.pdf |
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| format = [[PDF]] |
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| id = |
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| accessdate = 2006-06-30 |
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| doi = 10.1038/23856 |
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}} {{Dead link|date=September 2010|bot=H3llBot}} |
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</ref> At the beginning of his second paragraph on the peppered moths, Majerus emphasizes that the wealth of additional data obtained since Kettlewell's initial predation papers does not undermine the basic qualitative deductions from that work and that differential bird predation of the dark and light moths in habitats affected by industrial pollution to different degrees ([[directional selection]]) "is the primary influence of the evolution of melanism in the peppered moth".<ref name=ioo/><ref name="majerus1998">Majerus, M.E.N. (1998) ''[[Melanism: Evolution in Action]]''. Oxford University Press, New York.</ref> Coyne had erred in his statement that only two peppered moths had been found on tree trunks, as the book gives the resting positions of 47 peppered moths Majerus had found in the wild between 1964 and 1996; twelve were on tree trunks (six exposed, six unexposed), twenty were at the trunk/branch joint, and fifteen resting on branches.<ref name=ioo/> Majerus found that the review did not reflect the factual content of the book or his own views,<ref name=pmbw2>{{cite web |url=http://www.asa3.org/archive/evolution/199903/0312.html |title=Evolution - March 1999: Peppered Moths - in black and white (part 2 of 2) |accessdate=2007-08-26 |author=Donald Frack |date=30 March 1999}}</ref> and cites an assessment by the [[entomology|entomologist]] Donald Frack that there was essentially no resemblance between the book and Coyne's review,<ref name="dd">{{cite web |url=http://www.gen.cam.ac.uk/Research/Majerus/Darwiniandisciple.doc |title=The Peppered moth: decline of a Darwinian disciple |accessdate=2007-09-10 |author=Michael Majerus |authorlink=Michael Majerus |coauthors= |year=2004 |format=.doc |work= |publisher= |pages= |language= |archiveurl = http://web.archive.org/web/20070926131158/http://www.gen.cam.ac.uk/Research/Majerus/Darwiniandisciple.doc <!-- Bot retrieved archive --> |archivedate = 2007-09-26 |quote=}}</ref> which appeared to be a summary of the Sargent et al. paper rather than Majerus's book.<ref>{{cite web |url=http://www.asa3.org/archive/evolution/199903/0314.html |title=Evolution - March 1999: RE: Peppered Moths - in black and white (part 1 of 2) |accessdate=2007-08-26 |author=Donald Frack |date=30 March 1999}}</ref> |
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The review was subsequently picked up by journalist Robert Matthews, who wrote an article for ''[[Sunday Telegraph|The Sunday Telegraph]]'', March 14th, 1999, claiming that "Evolution experts are quietly admitting that one of their most cherished examples of Darwin's theory, the rise and fall of the peppered moth, is based on a series of scientific blunders. Experiments using the moth in the Fifties and long believed to prove the truth of natural selection are now thought to be worthless, having been designed to come up with the 'right' answer". Majerus regarded this view as surprising, and not one that would be shared by those involved in the field. He noted numerous scientific inaccuracies, misquotations and misrepresentations in the article, but thought this was common in press reports.<ref name="dd"/> He stated that he had spoken to Matthews for over half an hour and had to explain many details as Matthews hadn't read the book, but "Even then, he got nearly everything wrong."<ref name=pmbw2/> |
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The story was taken up by creationists, and at a seminar presenting the [[wedge strategy]] on March 13th, 1999, the leading intelligent design proponent [[Phillip E. Johnson]] said that the moths "do not sit on tree trunks", "moths had to be glued to the trunks" for pictures and that the experiments were "fraudulent" and a "scam."<ref>{{cite web |url=http://www.asa3.org/archive/evolution/199904/0201.html |title=Evolution - April 1999: Peppered Moths and Creationists |accessdate=2007-08-26 |author=Donald Frack |date=16 April 1999 }}</ref> This led Frack to exchanges with intelligent design proponent [[Jonathan Wells (intelligent design advocate)|Jonathan Wells]], who <!--bit off topic: studied biology as part of his life goal of "destroying Darwinism".<ref>[[Jonathan Wells (intelligent design advocate)|Wells, Jonathan]]. "[http://www.tparents.org/library/unification/talks/wells/DARWIN.htm Darwinism: Why I Went for a Second PhD]"</ref>--> conceded that Majerus listed six moths on exposed tree trunks (out of 47), but argued that this was "an insignificant proportion".<ref>{{cite web |url=http://www.asa3.org/archive/evolution/199904/0207.html |title=Evolution - April 1999: RE: My last word |accessdate=2007-08-26|author=Donald Frack |date=16 April 1999 }}</ref> Wells wrote an essay on the subject, a shortened version of which appeared in ''[[The Scientist]]'' of May 24th, 1999, claiming that "In 25 years of fieldwork, C.A. Clarke and his colleagues found only one peppered moth on a tree trunk", and concluding that "The fact that peppered moths do not normally rest on tree trunks invalidates Kettlewell's experiments".<ref name="wells1999">Wells, J. (1999). "[http://www.arn.org/docs/wells/jw_pepmothshort.htm Second Thoughts about Peppered Moths; This classical story of evolution by natural selection needs revising]." ''The Scientist'' '''13''', 11 (p. 13, May 24, 1999)</ref> In 2000 he wrote ''[[Icons of Evolution]]: Why much of what we Teach About Evolution is Wrong'', which claims "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."<ref name="wells2000">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 [http://www.iconsofevolution.com/ Iconsofevolution.com])</ref> The arguments put by Wells have been dismissed by Majerus, Cook and peppered moth researcher [[Bruce Grant]] who describes Wells as distorting the picture by selectively omitting or scrambling references in a way that is dishonest.<ref name=ioo/> |
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On November 27th, 2000, the school board of [[Pratt County, Kansas]] continued efforts to favor intelligent design teaching by requiring the use of specific resources.<ref>{{cite web |url=http://ncse.com/rncse/20/4/intelligent-design-pratt-county-kansas |title=Intelligent Design in Pratt County, Kansas |accessdate=2007-08-28 |author=Molleen Matsumura |work= |publisher= [[National Center Science Education]]}}</ref> These included the article by Jerry Coyne, who wrote to object strongly to this creationist misrepresentation of his critical re-evaluation, emphasising that the moth story is a sound example of evolution produced by natural selection and stating that his call for additional research was only to resolve uncertainty regarding bird predation as the cause of the natural selection and evolutionary change. Bruce Grant also wrote to challenge allegations of fraud in the moth experiments based on misrepresentations by Wells.<ref>{{cite web |url=http://www.indiana.edu/~ensiweb/lessons/icon.cr.html |title=Icons of Evolution |accessdate=2007-08-28 |author= |authorlink= |coauthors= |date= |work=Evolution and the Nature of Science Institutes for High School Biology Teachers: Resources |publisher=Department of Biology, Indiana University }}</ref> In 2002, Judith Hooper's ''[[Of Moths and Men]]'' added to the accusations of scientific fraud. She accused Kettlewell of manipulating his data to prove his hypothesis.<ref name=test>[http://www.newscientist.com/channel/opinion/mg19626330.500-reclaiming-the-peppered-moth-for-science.html Reclaiming the peppered moth for science] New Scientist, 08 December 2007, page 46-49</ref> The book received strong criticism from the scientific press (e.g., Coyne, B.C. Clarke, Grant).<ref name="coyne2002">Coyne, J.A. (2002). (Review of ''[[Of Moths and Men]]'')</ref><ref>{{cite web |url=http://www.talkreason.org/articles/moonshine.cfm |title=Talk Reason: arguments against creationism, intelligent design, and religious apologetics |accessdate=2007-08-28 |author=Matt Young |date= |work=Moonshine: Why the Peppered Moth remains an Icon of Evolution |publisher=Department of Physics, Colorado School of Mines |pages= |language= |archiveurl= |archivedate= |quote=}}</ref> Majerus described it as "littered with errors, misrepresentations, misinterpretations and falsehoods".<ref name="dd"/> |
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== See also == |
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*[[Polymorphism (biology)|Polymorphism]] |
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== References == |
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{{reflist|2}} |
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==External links== |
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{{commons|Biston betularia}} |
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* [[Bruce Grant]] has written several papers on melanism in the peppered moth which are listed on [http://bsgran.people.wm.edu his home page]. |
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* Online lecture: [http://www.streaming.mmu.ac.uk/cook/ "The rise and fall of the melanic Peppered Moth"] presented by Laurence Cook. |
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* {{cite web |url=http://www.talkreason.org/articles/moonshine.cfm |title=Moonshine: Why the Peppered Moth remains an Icon of Evolution |accessdate=2009-01-03 |author=Matt Young |date= |work= |publisher=Department of Physics, Colorado School of Mines |pages= |language= |archiveurl= |archivedate= |quote=}} |
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* [http://www.gen.cam.ac.uk/research/personal/majerus/Darwiniandisciple.pdf 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. |
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* [http://www.gen.cam.ac.uk/research/personal/majerus/Swedentalk220807.pdf 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 [http://www.gen.cam.ac.uk/research/personal/majerus/SwedenPepperedmoth2007Ppt.pdf also available]. |
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*{{cite web |url=http://www.springerlink.com/content/h7n4r6h026q1u6hk/fulltext.html |title=Industrial Melanism in the Peppered Moth, ''Biston betularia'': An Excellent Teaching Example of Darwinian Evolution in Action |author=Michael E. N. Majerus |authorlink=Michael Majerus |date= 6 December 2008 |work=10.1007/s12052-008-0107-y |publisher=Springer Science+Business Media, LLC 2008 |quote=Accusations of data fudging and scientific fraud in the case are found to be vacuous. |accessdate=2009-01-13}} |
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* On 19 June 2009, [http://www.telegraph.co.uk Telegraph.co.uk] published [http://www.telegraph.co.uk/earth/wildlife/5577724/Moth-turns-from-black-to-white-as-Britains-polluted-skies-change-colour.html an article] on this evolutionary phenomenon and implored UK readers to visit the [http://www.mothscount.org 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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{{pepperedmoth}} |
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[[Category:Peppered moth|Evolution]] |
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[[Category:Biology experiments]] |
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[[Category:Evolution by taxon]] |
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[[Category:Selection]] |
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[[de:Industriemelanismus]] |
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[[id:Evolusi biston betularia]] |
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[[ja:工業暗化]] |
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[[simple:Peppered moth evolution]] |
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[[fi:Teollisuusmelanismi]] |
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Revision as of 16:16, 9 February 2012
This is fake. I like Peanut Butter.