Evolution: Difference between revisions

From Citizendium
Jump to navigation Jump to search
imported>DavidGoodman
(needs dividing, see talk)
 
imported>Anthony.Sebastian
mNo edit summary
 
(86 intermediate revisions by 15 users not shown)
Line 1: Line 1:
{{otheruses4|evolution in biology}}
{{subpages}}
[[Image:Phylogenetic_tree.svg|thumb|350px|A hypothetical [[phylogenetic tree]] of all extant organisms, based on 16S [[non-coding RNA|rRNA]] [[gene]] sequence data, showing the evolutionary history of the [[three-domain system|three domains of life]], [[bacteria]], [[archaea]] and [[eukaryote]]s. Originally proposed by [[Carl Woese]].]]


In [[biology]], '''evolution''' is the change in the [[heritability|heritable]] [[trait (biology)|traits]] of a [[population genetics|population]] over successive generations, as determined by shifts in the [[allele frequency|allele frequencies]] of [[gene]]s. Over time, this process can result in [[speciation]], the development of new [[species]] from existing ones. All contemporary [[organism]]s are related to each other through [[common descent]], the products of cumulative evolutionary changes over billions of years. Evolution is the source of the vast [[biodiversity]] on [[Earth]], including the many [[extinction|extinct]] species attested in the [[fossil record]].<ref name=Futuyma>{{cite book | last = Futuyma | first = Douglas J. | authorlink = Douglas J. Futuyma | year = 2005 | title = Evolution | publisher = Sinauer Associates, Inc | location = [[Sunderland, Massachusetts|Sunderland]], [[Massachusetts]] | id = ISBN 0-87893-187-2}}</ref><ref>{{cite book | last = Gould | first = Stephen J. | authorlink = Stephen Gould  | year = 2002 | title = The Structure of Evolutionary Theory | publisher = Belknap Press | id = ISBN 0-674-00613-5}}</ref>
In [[biology]], the concept of '''evolution''' applies to a natural process whereby a distinct population of living systems, a biological species, for example, changes in its biological characteristics over generations, often with many new and diverse types of living systems emerging in the process. That application of the word 'evolution' to living systems does not exhaust the biological systems the word applies to; it includes transgenerational molecular evolution and [[Biolinguistics|language evolution]], for example.


The basic mechanisms that produce evolutionary change are [[natural selection]] (which includes [[ecological selection|ecological]], [[sexual selection|sexual]], and [[kin selection|kin]] selection) and [[genetic drift]]; these two mechanisms act on the genetic variation created by [[mutation]], [[genetic recombination]] and [[gene flow]]. Natural selection is the process by which individual organisms with favorable traits are more likely to survive and [[biological reproduction|reproduce]]. If those traits are heritable, they are passed to succeeding generations, with the result that beneficial heritable traits become more common in the next generation.<ref>{{cite journal | author = Lande, R. | coauthors = Arnold, S.J. | year = 1983 | title = The measurement of selection on correlated characters | journal = [[Evolution (journal)|Evolution]] | volume = 37 | pages = 1210&ndash;1226}}</ref><ref name=Futuyma/><ref>{{cite journal | author = Haldane, J.B.S. | year = 1953 | title = The measurement of natural selection | journal = Proceedings of the 9th International Congress of Genetics | volume = 1 | pages = 480&ndash;487}}</ref> Given enough time, this passive process can result in varied [[adaptation]]s to changing environmental conditions.<ref name="understandingevolution">{{cite web | url = http://evolution.berkeley.edu/evolibrary/article/0_0_0/evo_14 | title = Mechanisms: the processes of evolution | accessdate = 2006-07-14 | work = Understanding Evolution | publisher = [[University of California, Berkeley]]}}</ref>
In his 1998 edition of ''Evolutionary Biology'', evolutionary biologist Douglas Futuyma defined 'evolution' thus:
{|align="center" style="width:90%;font-size:98%;"
|
<font face="Gill Sans MT">"In its broadest sense, ''evolution'' simply means "change"...Usually..we don't apply the term to the changes an individual entity undergoes during its lifetime [i.e., its '[[ontogeny]]']. Rather, an evolving system is ordinarily one in which there is descent of entities [viz., populations of similar entities], one generation from another, over time, and in which characteristics of the entities differ across generations. Thus evolution in a broad sense is ''descent with modification'', and often ''with diversification''."</font><ref name=futuymaevobiol98>Futuyma DJ. (1998) Evolutionary Biology 3rd ed. Sinauer Associates, Inc., Sunderland MA ISBN 0-87893-189-9</ref>
|}
 
Expanding in his 2009 edition of ''Evolution'', he writes:
{|align="center" style="width:90%;font-size:98%;"
|
<font face="Gill Sans MT">"It [‘evolution’] is sometimes used to describe changes in individual objects such as stars. '''Biological''' (or '''organic'') '''evolution''', however, is ''change in the properties of groups of organisms over the course of generations''. The development, or ONTOGENY, of an individual organism is not consid¬ered evolution: individual organisms do not evolve. Groups of organisms, which we may call '''populations''', undergo ''descent with modification''. Populations may become subdivided, so that several populations are derived from a ''common ancestral population''. If different changes transpire in the several populations, the populations ''diverge''.”</font><ref name=futuevo2009>Futuyma DJ. (2009) ''Evolution''. 2nd ed. Sunderland, MA: Sinauer Associates. ISBN 978-0-87893-223-8.</ref>
|}


The modern understanding of evolution is based on the theory of natural selection, which was first set out in a joint 1858 paper by [[Charles Darwin]] and [[Alfred Russel Wallace]] and popularized in Darwin's 1859 book ''[[The Origin of Species]]''. In the 1930s, Darwinian natural selection was combined with the theory of [[Gregor Mendel|Mendelian]] [[heredity]] to form the [[modern evolutionary synthesis]], also known as "Neo-[[Darwinism]]". The modern synthesis describes evolution as a change in the frequency of [[allele]]s within a population from one generation to the next.<ref name="understandingevolution"/> With its explanatory and [[predictive power]], this theory has become the central organizing principle of modern biology, relating directly to topics such as the origin of [[antibiotic resistance]] in bacteria, [[eusociality]] in insects, and the staggering [[biodiversity]] of Earth's [[ecosystem]].  
In one of the specific forms the process of evolution unrolls, a closely related group of organisms, referred to as a 'population', undergoes modifications of its characteristics&mdash;traits, or so-called [[phenotype]]s&mdash;over a period of generations in a manner that subdivides a parent population into distinctly differing descendant populations, a process in which a common ancestral population gives rise to diverse populations&mdash;the process of change referred to as 'descent with modification'.  
{{TOC|left}}
During the 1800s, before mid-century, two factors in combination had made biological evolution plausible and likely to open-minded leading scientists, such as the eminent geologist, Charles Lyell<ref>[http://books.google.co.uk/books?id=mFkOAAAAQAAJ&pg=PA10&ots=GqTXDf8Ywe&dq=charles+Lyell&output=text#c_top ''Principles of Geology'']</ref>, and biologist [[Alfred Russel Wallace]]. The first: Evidence from strategraphic geology suggested a much older age of the earth than the approximately 4,000 years biblical scholars had determined for the Christian orthodox view, important evidence because of a constraining power of Christianity on the acceptance of scientific ideas that contradicted accepted interpretation of the Christian bible. No less important, the much older age of the earth would have extended greatly a time for biological evolution to occur. The second: Evidence from paleontology indicated the existence of unfamiliar species from the time of the old earth, in the old geographic strata, now extinct.


Although there is overwhelming [[scientific consensus]] supporting the validity of evolution, because of its potential implications for the origins of humankind, evolutionary theory has been at the center of many [[creation-evolution controversy|social and religious controversies]] since its inception.  
Thus, biological evolution was plausible and considered likely in science by the mid-1800s because the earth appeared very old and because species existed in the old earth, but not the contemporary species, extinct forms from which contemporary species might have descended by transformation. By mid-century the time was ripe for theories to explain how a descendant evolution might occur, and for a few, ripe for strictly naturalistic theories.


{{evolution3}}
The word 'evolution' derives from the Latin word ''evolvere'', which translates to 'to unfold', or 'to unroll', as if to unroll an ancient scroll and display the intellectual potentialities hidden within, or to unfold the map of time-development of biological diversity.<ref name=futuyma2005>Futuyma DJ. (2005) ''Evolution''. 1st ed. Sunderland, Mass.: Sinauer Associates. ISBN 0878931872. | [http://www.sinauer.com/evolution/sample/ Downloadable Chapter 15 2nd.  ed. "Sex and Reproductive Success"].</ref> Before the word 'evolution' was introduced in biology to refer to the process of transgenerational changes in species, the process was referred to as '[[transmutation]]', or 'transformation', the concepts, '[[transmutationism]]' and 'transformationism'.<ref>Wilkins J. (1997) [http://www.biologie.uni-hamburg.de/b-online/e36_talk/precursors/precurs2.html Darwin's Precursors and Influences: Introduction]</ref>


== Study of evolution ==
As mentioned above, 'evolution' applies not only to biological evolution, the topic of the present article, but also to other domains of nature that change in a descendant way similar to that of biological evolution, such as the evolution of the universe itself.<ref name=smolin2008>Smolin L. (2008) ''The Life of the Cosmos''. Oxford University Press. ISBN 9780199216802. | [http://books.google.com/books?id=zdC0GKsgnEwC&dq=the+life+of+the+cosmos&source=gbs_navlinks_s Google Books preview].</ref>
{{main|Evolutionary biology}}


=== History of evolutionary thought ===
<!--Save this paragraph for mechanism-related technicalities below:
{{main|History of evolutionary thought}}
Technically, evolution involves change in the [[heritability|heritable]]<ref name=visscher2008>Visscher PM, Hill WG, Wray NR. (2008) [http://dx.doi.org/10.1038/nrg2322 Heritability in the genomics era &mdash; concepts and misconceptions] ''Nature Reviews Genetics'' 9:255-266.
[[Image:Charles Darwin.jpg|frame|left|[[Charles Darwin]] in 1854, five years before publishing ''[[The Origin of Species]]''.]]
*<font face="Gill Sans MT"><u>From the Abstract:</u>&nbsp;Heritability allows a comparison of the relative importance of genes and environment to the variation of traits within and across populations. The concept of heritability and its definition as an estimable, dimensionless population parameter was introduced by Sewall Wright and Ronald Fisher nearly a century ago. Despite continuous misunderstandings and controversies over its use and application, heritability remains key to the response to selection in evolutionary biology and agriculture, and to the prediction of disease risk in medicine.</font></ref> [[trait (biology)|traits]] of a [[population genetics|population]] over successive generations, as determined by shifts in the [[allele frequency|allele frequencies]] of [[gene]]s. Over time, this process can result in [[speciation]], the development of new [[species]] from existing ones. A main assumption in evolutionary theory is that contemporary [[organism]]s are related to each other through [[common descent]] from a single, or small number, of distantly common ancestors - the first life forms on earth that succeeded in giving rise to viable offspring.-->


The idea of biological evolution has existed since ancient times, notably among Greek philosophers such as [[Anaximander]] and [[Epicurus]] and Indian philosophers such as [[Patañjali]]. However, scientific theories of evolution were not proposed until the 18th and 19th centuries, by scientists such as [[Jean-Baptiste Lamarck]] and [[Charles Darwin]].
Among the many ways biologists recognize the explanatory power of the concept of evolution, they can account for the vast [[biodiversity]] of living systems on Earth, including the many [[extinction|extinct]] species attested in the [[fossil record]]. An evolutionary perspective can lead to [[Evolutionary medicine|insights for preventing, diagnosing and treating human diseases]].


The [[transmutation of species]] was accepted by many scientists before 1859, but Charles Darwin's ''[[The Origin of Species|On The Origin of Species by Means of Natural Selection]]'' provided the first convincing exposition<ref>In the years after Darwin's publication and fame, numerous "predecessors" to natural selection were discovered, such as [[William Charles Wells]] and [[Patrick Matthew]], who had published unelaborated and undeveloped versions of similar theories earlier to little or no attention. Historians acknowledge that Darwin was the first to develop the theory rigorously and developed it independently. On Matthew, one historian of evolution has written that he "did suggest a basic idea of selection, but he did nothing to develop it; and he published it in the appendix to a book on the raising of trees for shipbuilding. No one took him seriously, and he played no role in the emergence of Darwinism. Simple priority is not enough to earn a thinker a place in the history of science: one has to develop the idea and convince others of its value to make a real contribution. Darwin's notebooks confirm that he drew no inspiration from Matthew or any of the other alleged precursors." {{cite book | last = Bowler | first = Peter J. | authorlink = Peter J. Bowler | year = 2003 | title = Evolution: The History of an Idea | publisher = University of California Press | location = Berkeley| pages =158}}</ref> of a mechanism by which evolutionary change could occur: [[natural selection]]. After many years of working in private on his theory, Darwin was motivated to publish his work on evolution when he received a letter from Alfred Russel Wallace in which Wallace revealed his own, independent discovery of natural selection. Accordingly, Wallace is sometimes given shared credit for originating the theory.<ref>{{cite book | last = Bowler | first = Peter J. | authorlink = Peter J. Bowler | year = 2003 | title = Evolution: The History of an Idea | publisher = University of California Press | location = Berkeley}}</ref>
Most biologists endorse the proposition of the 20th century's pioneering evolutionary biologist, [[Theodosius Dobzhansky]], ''videlicet'': "Nothing in biology makes sense except in the light of evolution."<ref name="Dobzhansky1973">{{cite journal |author=Dobzhansky T |title=Nothing in biology makes sense except in the light of evolution | journal=Am Biol Teach |pages=125-9 |year=1973 |url=http://www.godslasteraar.org/assets/ebooks/Dobzhansky_Nothing_in_Biology_Makes_Sense_Except_in_the_Light_of_Evolution.pdf }}</ref>&nbsp;&nbsp;For ''evolutionary'' biologists, not surprisingly, [[Evolution|evolution ]] takes center stage, in particular as "the unifying theory of biology".<ref name=futuymaevobiol98/>&nbsp;<ref>[http://www.talkorigins.org/faqs/faq-intro-to-biology.html Introduction to Evolutionary Biology]</ref>


The publication of Darwin's book sparked a great deal of scientific and social debate. Darwin's work relied on many different fields of scientific inquiry for its evidence, and as a consequence debates over the theory took place in many different arenas. Although the occurrence of biological evolution of some sort came to be widely accepted by scientists, Darwin's specific ideas about evolution&mdash;that it occurred gradually, through natural selection&mdash;were actively attacked and contested. Additionally, while Darwin was able to observe variation, and infer natural selection and thereby adaptation, he was unable to explain ''how'' variation might arise, or be altered over generations. Darwin's proposal of a [[heredity|hereditary]] mechanism ([[pangenesis]]) lacked evidence and was ultimately rejected,<ref>[http://scienceclassics.nas.edu/moore/chap01.html Darwin’s Theory of Pangenesis]</ref> being replaced by [[genetics]].
Biologists distinguish between the factuality of evolution and theories of its [[Natural selection|mechanism]].  


From the end of the 19th century through the early 20th century, forms of neo-Lamarckism, "progressive" evolution ([[orthogenesis]]), and an evolution which worked by "jumps" ([[saltation (biology)|saltationism]], as opposed to [[phyletic gradualism|gradualism]]) became popular, although a form of neo-Darwinism, led by [[August Weismann]], also enjoyed some minor success. The biometric school of evolutionary theory, resulting from the work of Darwin's cousin, [[Francis Galton]], emerged as well, using statistical approaches to biology which emphasized gradualism and some aspects of natural selection.<ref name="mendelianrevolution">{{cite book | last = Bowler | first = Peter J. | authorlink = Peter J. Bowler | year = 1989 | title = The Mendelian Revolution: The Emergence of Hereditarian Concepts in Modern Science and Society | publisher = John Hopkins University Press | location = Baltimore}}</ref>
Although there is overwhelming [[scientific consensus]] supporting the reality of biological evolution, because of its potential implications for the origins of humankind, evolutionary theory has been at the center of many [[creation-evolution controversy|social and religious controversies]] since its inception.
<!--Save  this paragraph for a mechanism section below:
The basic mechanisms that produce evolutionary change are [[natural selection]] (also referred to as 'survival of the fittest) and [[genetic drift]].  Natural selection was first described in 1858 in two papers, jointly published, one by [[Charles Darwin]] and the other by [[Alfred Russel Wallace]], and then popularized in Darwin's 1859 book ''[[The Origin of Species]]''.  It is the process by which individual organisms with favorable traits are more likely to survive and [[biological reproduction|reproduce]]. If those traits are heritable, they are passed to succeeding generations, with the result that beneficial heritable traits become more common in the next generation.<ref>{{cite journal | author = Lande, R. | coauthors = Arnold, S.J. | year = 1983 | title = The measurement of selection on correlated characters | journal = [[Evolution (journal)|Evolution]] | volume = 37 | pages = 1210&ndash;1226}}</ref><ref>{{cite journal | author = Haldane JBS | year = 1953 | title = The measurement of natural selection | journal = Proceedings of the 9th International Congress of Genetics | volume = 1 | pages = 480-7}}</ref> Given enough time, this passive process can result in varied [[adaptation]]s to changing environmental conditions.<ref name="understandingevolution">{{cite web | url = http://evolution.berkeley.edu/evolibrary/article/0_0_0/evo_14 | title = Mechanisms: the processes of evolution | accessdate = 2006-07-14 | work = Understanding Evolution | publisher = [[University of California, Berkeley]]}}</ref>-->


[[Image:Mendel.png|frame|right|[[Gregor Mendel]]'s work on the [[Mendelian inheritance|inheritance]] of traits in pea plants laid the foundation for [[genetics]].]]
=== Brief history of the idea of evolution ===
The concept that plants and animals change form over generations has existed since ancient times, notably among Greek philosophers such as [[Anaximander]]<ref name=hxecoiep>[http://www.iep.utm.edu/evolutio/ History of Evolution]. Internet Encyclopedia of Philosophy.</ref>
and [[Epicurus]]<ref name=konstan>Konstan D. (2009) "Epicurus", [http://plato.stanford.edu/archives/spr2009/entries/epicurus/ ''The Stanford Encyclopedia of Philosophy (Spring 2009 Edition)'', Edward N. Zalta (ed.)
*<font face="Gill Sans MT">"The philosophy of Epicurus (341–270 B.C.) was a complete and interdependent system, involving a view of the goal of human life (happiness, resulting from absence of physical pain and mental disturbance), an empiricist theory of knowledge (sensations, including the perception of pleasure and pain, are infallible criteria), a description of nature based on atomistic materialism, and <i>a naturalistic account of evolution, from the formation of the world to the emergence of human societies.</i>" [emphasis added]</font></ref>&nbsp;<ref name=evoepiwik>[http://wiki.epicurus.info/Evolution Evolution - Epicurus Wiki]</ref> and Indian philosophers such as [[Patañjali]]
{|align="center" style="width:90%;font-size:98%;"
|
<font face="Gill Sans MT"> Evolution is not so much a modern discovery as some of its advocates would have us believe. It made its appearance early in Greek philosophy, and maintained its position more or less, with the most diverse modifications, and frequently confused with the idea of emanation, until the close of ancient thought. The Greeks had, it is true, no term exactly equivalent to ” evolution”; but when Thales asserts that all things originated from water; when Anaximenes calls air the principle of all things, regarding the subsequent process as a thinning or thickening, they must have considered individual beings and the phenomenal world as, a result of evolution, even if they did not carry the process out in detail. Anaximander is often regarded as a precursor of the modem theory of development. He deduces living beings, in a gradual development, from moisture under the influence of warmth, and suggests the view that men originated from animals of another sort, since if they had come into existence as human beings, needing fostering care for a long time, they would not have been able to maintain their existence. In Empedocles, as in Epicurus and Lucretius, who follow in Hs [sic] footsteps, there are rudimentary suggestions of the Darwinian theory in its broader sense; and here too, as with Darwin, the mechanical principle comes in; the process is adapted to a certain end by a sort of natural selection, without regarding nature as deliberately forming its results for these ends.</font> <ref name=hxecoiep>[http://www.iep.utm.edu/evolutio/ History of Evolution]. Internet Encyclopedia of Philosophy.</ref>
|}
The ancient Greeks started the naturalists and devotees of natural history, as well as the world of science known as natural philosophy, on the path of thinking about change in the animate and inanimate world and the implications of those thoughts on the changes in the recent past and on future change. By the mid-1400s ideas of evolution began fast growing in abundance. Them concept of evolution responded to the fuel of expanding knowledge of fossilized living things, and the recognition of Earth's history, its long time of existence, coming up with the idea that the available time favored evolution of present living things from ancestors that looked different in their fossilized  remains.


Darwin's lack of a hereditary mechanism is often seen today as a major stumbling block in the historical acceptance of his theory, but in his time it was not a pressing issue, as questions of the development of an organism were seen as more important than questions of the transmission of hereditary traits; Darwin and other biologists of his day thought that the answers to heredity would be found in [[embryology]] rather than in breeding experiments. Work on plant hybridity by a contemporary of Darwin's, an [[Augustinians|Augustinian]] monk in [[Bohemia]] named [[Gregor Mendel]], revealed that certain traits in [[pea]]s occurred in discrete forms (that is, they were either one distinct trait or another,  such as "round" or "wrinkled") and were inherited in a well-defined and predictable manner. Mendel's [[Mendelian inheritance#Mendel.27s law of segregation|Law of Segregation]] and [[Mendelian inheritance#Mendel.27s law of independent assortment|Law of Independent Assortment]] would eventually become key theories in the development of [[genetics]], but in Darwin's time their significance was not seen (even by Mendel himself).<ref name="mendelianrevolution"/>
== Mechanisms that mediate evolution ==
[[Jean-Baptiste Lamarck]] theorized that living things passed on their traits to offspring, but his theories differed from modern evolutionary theory in that he believed that acquired traits were passed on. Thus, Lamarckian evolution posits that the giraffe acquired its long neck from stretching to reach higher foliage. An individual giraffes lengthened its neck from a lifetime of stretching, and passed on the longer neck to its offspring, which themselves stretched to reach even-higher branches, creating a cycle that concluded with the long necks of today's giraffes.<ref>[http://evolution.berkeley.edu/evosite/history/evol_happens.shtml Early Concepts of Evolution: Jean Baptiste Lamarck]</ref>


When Mendel's work was "rediscovered" in 1901, it was initially interpreted as supporting an anti-Darwinian "jumping" form of evolution. The convinced Mendelians, such as [[William Bateson]] and [[Charles Benedict Davenport]], and biometricians, such as [[Walter Frank Raphael Weldon]] and [[Karl Pearson]], became embroiled in a bitter debate, with Mendelians charging that the biometricians did not understand biology, and biometricians arguing that most biological traits exhibited continuous variation rather than the "jumps" expected by the early Mendelian theory.<ref name="diftraits">It is now recognized that the Mendelians were investigating ''Mendelian traits'', those traits where existing variation is controlled by one gene and therefore is discrete, and the biometricians were investigating ''complex traits'', where those traits were controlled by multiple genes, and the variation is therefore continuous.</ref> However, the simple version of the theory of early Mendelians soon gave way to the [[classical genetics]] of [[Thomas Hunt Morgan]] and his school, which thoroughly grounded and articulated the applications of Mendelian laws to biology. Eventually, it was shown that a rigorous statistical approach to Mendelism was reconcilable with the data of the biometricians by the work of statistician and population geneticist [[Ronald Fisher|R.A. Fisher]] in the 1930s. Following this, the work of population geneticists&mdash;notably [[Sewall Wright]] and [[J. B. S. Haldane]]&mdash;and zoologists in the 1930s and 1940s synthesized Darwinian evolution with genetics, creating the [[modern evolutionary synthesis]].<ref name="mendelianrevolution"/> Genes were then still theoretical entities, and many paleontologists and embryologists were inclined to dismiss them as being of no, or minor, importance, but subsequent advancements have made genetics a key aspect of evolutionary biology.<ref>Resynthesizing evolutionary and developmental biology. Gilbert SF, Opitz JM, Raff RA. Developmental Biology 1996 Feb 1;173(2):357-72</ref>
The [[transmutation of species]] was accepted by many scientists before 1859, but Charles Darwin's ''[[The Origin of Species|On The Origin of Species by Means of Natural Selection]]'' provided the first convincing exposition<ref>In the years after Darwin's publication, numerous "predecessors" to natural selection were discovered, such as [[William Charles Wells]] and [[Patrick Matthew]], who had published unelaborated and undeveloped versions of similar theories earlier to little or no attention. Darwin was the first to develop the theory rigorously and developed it independently. On Matthew, one historian of evolution has written that he "did suggest a basic idea of selection, but he did nothing to develop it; and he published it in the appendix to a book on the raising of trees for shipbuilding. No one took him seriously, and he played no role in the emergence of Darwinism. Simple priority is not enough to earn a thinker a place in the history of science: one has to develop the idea and convince others of its value to make a real contribution. Darwin's notebooks confirm that he drew no inspiration from Matthew or any of the other alleged precursors." {{cite book | last = Bowler | first = PJ | authorlink = Peter J. Bowler | year = 2003 | title = Evolution: The History of an Idea | publisher = University of California Press | location = Berkeley| pages =158}}</ref> of a mechanism by which evolutionary change could occur: [[natural selection]]. After many years of working in private on his theory, Darwin was motivated to publish his work on evolution when he received a letter from Alfred Russell Wallace in which Wallace revealed his own, independent discovery of natural selection. Accordingly, Wallace is usually given shared credit for originating the theory.<ref name=bowler2003> Bowler,P.J. (2003) ''Evolution: The History of an Idea''. 3rd ed., completely rev. and expanded. Berkeley: University of California Press. ISBN 052236939. | [http://books.google.com/books?id=gJXmS49Q7r0C&source=gbs_navlinks_s Google Books preview]. $<font face=Gill Sans MT">From chapter 1: "Much can be gained from the lessons of history, especially an awareness of the dangers of oversimplification. Before we can undertake such a study, however, we must establish a framework for understanding the rise of modern evolutionism. We must identify the full range of issues over which evolutionism has challenged the traditional worldview. We must appreciate how aspects of the old view can be modernized to fit in with certain kinds of evolutionary thought. The basic idea of evolution can be developed in many different ways, each with its own broader implications. We must also look more closely at the problems facing the historian who has to seek a balance between the conventional image of science as the objective evaluation of factual data and the suspicion of many critics that scientific theories are value-laden contributions to philosophical and ideological debates."</font></ref>


The most significant recent developments in evolutionary biology have been the improved understanding of and advances in [[genetics]].<ref>{{cite news | author = Rincon, Paul | url = http://news.bbc.co.uk/1/hi/sci/tech/4552466.stm | title = Evolution takes science honours | publisher = [[BBC News]] | date = 2005 | accessdate = 2006-07-16}} According to the [[BBC]], [[Colin Norman]], news editor of [[Science (journal)|Science]], said "[S]cientists tend to take for granted that evolution underpins modern biology [...] Evolution is not just something that scientists study as an esoteric enterprise. It has very important implications for public health and for our understanding of who we are" and Dr. Mike Ritchie, of the school of biology at the University of St Andrews, UK said "The big recent development in evolutionary biology has obviously been the improved resolution in our understanding of genetics. Where people have found a gene they think is involved in speciation, I can now go and look how it has evolved in 12 different species of fly, because we've got the genomes of all these species available on the web."</ref> In the 1940s, following up on [[Griffith's experiment]], [[Oswald Avery|Avery]], [[Colin MacLeod|MacLeod]] and [[Maclyn McCarty|McCarty]] definitively identified [[DNA]] (deoxyribonucleic acid) as the "transforming principle" responsible for transmitting genetic information. In 1953, [[Francis Crick]] and [[James D. Watson]] published their famous paper on the structure of DNA, based on the research of [[Rosalind Franklin]] and [[Maurice Wilkins]]. These developments ignited the era of [[molecular biology]] and transformed the understanding of evolution into a molecular process (see [[molecular evolution]]): the [[mutation]] of segments of DNA. [[George C. Williams]]' 1966 ''Adaptation and natural selection: A Critique of some Current Evolutionary Thought'' marked a departure from the idea of [[group selection]] towards the modern notion of the gene as the [[unit of selection]]. In the mid-1970s, [[Motoo Kimura]] formulated the [[neutral theory of molecular evolution]], firmly establishing the importance of [[genetic drift]] as a mechanism of evolution.
Darwin's book sparked a great deal of scientific and social debate. Darwin's work relied on many different fields of scientific inquiry for its evidence, and debates over the theory took place in many different arenas. Although the occurrence of biological evolution was generally accepted by scientists, Darwin's specific ideas about evolution&mdash;that it occurred gradually, through natural selection&mdash;were contested. Additionally, while Darwin was able to observe variation, and infer natural selection and thereby adaptation, he was unable to explain ''how'' variation might arise, or be altered over generations. Darwin's proposal of a [[heredity|hereditary]] mechanism ([[pangenesis]]) lacked evidence and was ultimately rejected,<ref>[http://scienceclassics.nas.edu/moore/chap01.html Darwin’s Theory of Pangenesis]</ref> being replaced by [[genetics]].


Debates over various aspects of how evolution occurs have continued. One prominent debate was over the theory of [[punctuated equilibrium]], proposed in 1972 by [[paleontology|paleontologists]] [[Niles Eldredge]] and [[Stephen Jay Gould]] to explain the paucity of gradual transitions between species in the fossil record, as well as the absence of change or stasis that is observed over significant intervals of time.
From the end of the 19th century through the early 20th century, forms of neo-Lamarckism, "progressive" evolution ([[orthogenesis]]), and an evolution which worked by "jumps" ([[saltation (biology)|saltationism]], as opposed to [[phyletic gradualism|gradualism]]) became popular, although a form of neo-Darwinism, led by [[August Weismann]], also enjoyed some minor success. The biometric school of evolutionary theory, resulting from the work of Darwin's cousin, [[Francis Galton]], emerged as well, using statistical approaches to biology which emphasized gradualism and some aspects of natural selection.<ref name="mendelianrevolution">{{cite book | last = Bowler | first = Peter J. | authorlink = Peter J. Bowler | year = 1989 | title = The Mendelian Revolution: The Emergence of Hereditarian Concepts in Modern Science and Society | publisher = John Hopkins University Press | location = Baltimore}}</ref>
 
Mendel's work was "rediscovered" in 1901.  Debates over various aspects of how evolution occurs have continued. One prominent debate was over the theory of [[punctuated equilibrium]], proposed in 1972 by [[paleontology|paleontologists]] [[Niles Eldredge]] and [[Stephen Jay Gould]] to explain the paucity of gradual transitions between species in the fossil record, as well as the absence of change or stasis that is observed over significant intervals of time.


===Academic disciplines===
===Academic disciplines===
Scholars in a number of academic disciplines continue to document examples of the theory of evolution, contributing to a deeper understanding of its underlying mechanisms. Every subdiscipline within [[biology]] both informs and is informed by knowledge of the details of evolution, such as in [[ecological genetics]], [[human evolution]], [[molecular evolution]], and [[phylogenetics]]. Areas of mathematics (such as [[bioinformatics]]), physics, chemistry and other fields all make important foundational contributions to the theory of evolution. Even disciplines as far removed as [[geology]] and [[sociology]] play a part, since the process of biological evolution has coincided in time and space with the development of both the Earth and human civilization.


[[Evolutionary biology]] is a subdiscipline of biology concerned with the origin and descent of [[species]], as well as their changes over time. It was originally an [[interdisciplinarity|interdisciplinary]] field including scientists from many traditional [[alpha taxonomy|taxonomically]]-oriented disciplines. For example, it generally includes scientists who may have a specialist training in particular organisms, such as [[mammalogy]], [[ornithology]], or [[herpetology]], but who use those organisms to answer general questions in evolution. Evolutionary biology as an [[academic discipline]] in its own right emerged as a result of the [[modern evolutionary synthesis]] in the 1930s and 1940s. It was not until the 1970s and 1980s, however, that a significant number of universities had departments that specifically included the term ''evolutionary biology'' in their titles.
[[Evolutionary biology]] is concerned with the origin and descent of [[species]], as well as their changes over time. It is an [[interdisciplinarity|interdisciplinary]] field including scientists from many traditional [[alpha taxonomy|taxonomically]]-oriented disciplines, including scientists with specialist training in particular organisms, such as [[mammalogy]], [[ornithology]], or [[herpetology]], but who use those organisms to answer general questions in evolution. Evolutionary biology as an [[academic discipline]] in its own right emerged as a result of the [[modern evolutionary synthesis]] in the 1930s and 1940s.  


[[Evolutionary developmental biology]] (informally, evo-devo) is a field of biology that compares the developmental processes of different animals in an attempt to determine the ancestral relationship between organisms and how developmental processes evolved. The discovery of [[gene]]s regulating development in model organisms allowed for comparisons to be made with genes and genetic networks of related organisms.
[[Evolutionary developmental biology]] (informally, evo-devo) is a field of biology that compares the developmental processes of different animals in an attempt to determine the ancestral relationship between organisms and how developmental processes evolved. The discovery of [[gene]]s regulating development in model organisms allowed for comparisons to be made with genes and genetic networks of related organisms.
Line 46: Line 68:
[[Physical anthropology]] emerged in the late 19th century as the study of human [[osteology]], and the fossilized skeletal remains of other [[Hominidae|hominid]]s. At that time, anthropologists debated whether their evidence supported Darwin's claims, because skeletal remains revealed temporal and spatial variation among hominids, but Darwin had not offered an explanation of the specific mechanisms that produce variation. With the recognition of Mendelian genetics and the rise of the modern synthesis, however, evolution became both the fundamental conceptual framework for, and the object of study of, physical anthropologists. In addition to studying skeletal remains, they began to study genetic variation among human populations ([[population genetics]]); thus, some physical anthropologists began calling themselves biological anthropologists.
[[Physical anthropology]] emerged in the late 19th century as the study of human [[osteology]], and the fossilized skeletal remains of other [[Hominidae|hominid]]s. At that time, anthropologists debated whether their evidence supported Darwin's claims, because skeletal remains revealed temporal and spatial variation among hominids, but Darwin had not offered an explanation of the specific mechanisms that produce variation. With the recognition of Mendelian genetics and the rise of the modern synthesis, however, evolution became both the fundamental conceptual framework for, and the object of study of, physical anthropologists. In addition to studying skeletal remains, they began to study genetic variation among human populations ([[population genetics]]); thus, some physical anthropologists began calling themselves biological anthropologists.


==Evidence of evolution==
===Fossil Record===
{{main|Evidence of evolution}}
[[Fossil]]s are critical evidence for estimating when various lineages originated. Since fossilization of an organism is an uncommon occurrence, usually requiring hard parts (like teeth, bone or pollen), the [[fossil record]] is traditionally thought to provide only sparse and intermittent information about ancestral lineages. Fossilization of organisms ''without'' hard body parts is rare, but does happen under unusual circumstance; very rapid burial, and both low oxygen environment and sparse microbial action<ref>{{cite journal| author =Schweitzer M.H. ''et al'' | year =2005| title = Soft-tissue vessels and cellular preservation in Tyrannosaurus rex| journal =Science | volume =307 | issue =5717 | pages =1952-1955}}</ref>.
Evolution has left numerous records which reveal the history of different species. [[fossil record|Fossils]], together with the [[comparative anatomy]] of present-day plants and animals, constitute the morphological, or [[anatomy|anatomical]], record. By comparing the anatomies of both modern and extinct species, [[paleontology|paleontologists]] can infer the lineages of those species. Important fossil evidence includes the connection of distinct classes of organisms by so-called "[[transitional fossil|transitional]]" species, such as the [[Archaeopteryx]], which provided early evidence for the link between [[dinosaur]]s and [[bird]]s,<ref>{{cite book | author = Feduccia, Alan | year = 1996 | title = The Origin and Evolution of Birds | publisher = Yale University Press | location = New Haven | id = ISBN 0-300-06460-8}}</ref> and the recently-discovered [[Tiktaalik]], which clarifies the development from [[fish]] to [[tetrapod|animals with four limbs]].<ref>{{cite journal | author = Daeschler, Edward B., Shubin, Neil H., & Jenkins Jr, Farish A. | year = 2006 | month = April | title = A Devonian tetrapod-like fish and the evolution of the tetrapod body plan | journal = [[Nature (journal)|Nature]] | volume = 440 | pages = 757&ndash;763 | doi = 10.1038/nature04639 | url = http://www.nature.com/nature/journal/v440/n7085/abs/nature04639.html | accessdate = 2006-07-14}}</ref>


The development of [[genetics|molecular genetics]], and particularly of DNA sequencing, has allowed biologists to study the record of evolution left in the organisms' genetic structures. The degree of similarity and difference in the DNA sequences of modern species allows geneticists to reconstruct their lineages. It is from DNA sequence comparisons that figures such as the 95% similarity between humans and chimpanzees are obtained.<ref>{{cite journal | author = Chimpanzee Sequencing and Analysis Consortium | year = 2005 | title = Initial sequence of the chimpanzee genome and comparison with the human genome | journal = [[Nature (journal)|Nature]] | volume = 437 | pages = 69&ndash;87}}</ref><ref>{{cite journal | author = Britten, R.J. | year = 2002 | title = Divergence between samples of chimpanzee and human DNA sequences is 5%, counting indels | journal = Proc Natl Acad Sci U S A | volume = 99 | pages = 13633&ndash;13635}}</ref>
The fossil record contains the earliest known examples of life itself, as well as the earliest occurrences of individual [[Lineage (evolution)|lineages]]. The first complex animals date from the [[early Cambrian]] period, approximately 520 million years ago. The records of individual species yield information regarding the patterns and rates of evolution, showing for example if species evolve into new species (speciation) gradually and incrementally, or in relatively brief intervals of geologic time. The fossil record is a document of large scale patterns and events in the history of life, many of which have influenced the evolutionary history of numerous [[Lineage (evolution)|lineages]]. [[Extinction event|Mass extinctions]] frequently resulted in the loss of entire groups of species, such as the non-avian dinosaurs, while leaving others relatively unscathed. Recently, molecular biologists have used the time since divergence of related lineages to calibrate the rate at which mutations accumulate, and at which the [[genomes]] of different lineages evolve.
 
Other evidence used to demonstrate evolutionary lineages includes the geographical distribution of species. For instance, [[monotreme]]s and most [[marsupial]]s are found only in [[Australia]], showing that their common ancestor with placental mammals lived before the submerging of the ancient [[land bridge]] between Australia and Asia.
 
Scientists correlate all of the above evidence, drawn from [[paleontology]], anatomy, genetics, and geography, with other information about the [[history of Earth]]. For instance, [[paleoclimatology]] attests to periodic [[ice age]]s during which the world's climate was much cooler, and these are often found to match up with the spread of species which are better-equipped to deal with the cold, such as the [[woolly mammoth]].
 
===Morphological evidence===
[[Image:Skelett vom Wal MK1888 ohne Text.gif|350px|thumb|Letter ''c'' in the picture indicates the undeveloped hind legs of a [[baleen whale]], [[vestigial structure|vestigial]] remnants of its terrestrial ancestors.]]
 
[[Fossil]]s are critical evidence for estimating when various lineages originated. Since fossilization of an organism is an uncommon occurrence, usually requiring hard parts (like teeth, bone or pollen), the [[fossil record]] is traditionally thought to provide only sparse and intermittent information about ancestral lineages. Fossilization of organisms without hard body parts is rare, but happens under unusual circumstances, such as rapid burial, low oxygen environments, or microbial action<ref>{{cite journal| author =Schweitzer M.H. ''et al'' | year =2005| title = Soft-tissue vessels and cellular preservation in Tyrannosaurus rex| journal =Science | volume =307 | issue =5717 | pages =1952-1955}}</ref>.
 
The fossil record provides several types of data important to the study of evolution. First, the fossil record contains the earliest known examples of life itself, as well as the earliest occurrences of individual [[Lineage (evolution)|lineages]]. For example, the first complex animals date from the [[early Cambrian]] period, approximately 520 million years ago. Second, the records of individual species yield information regarding the patterns and rates of evolution, showing for example if species evolve into new species (speciation) gradually and incrementally, or in relatively brief intervals of geologic time. Thirdly, the fossil record is a document of large scale patterns and events in the history of life, many of which have influenced the evolutionary history of numerous [[Lineage (evolution)|lineages]]. For example, [[Extinction event|mass extinctions]] frequently resulted in the loss of entire groups of species, such as the non-avian dinosaurs, while leaving others relatively unscathed. Recently, molecular biologists have used the time since divergence of related lineages to calibrate the rate at which mutations accumulate, and at which the [[genomes]] of different lineages evolve.


[[Phylogenetics]], the study of the ancestry of species, has revealed that structures with similar internal organization may perform divergent functions. [[Vertebrate]] limbs are a common example of such [[homology (biology)|homologous]] structures. The appendages on bat wings, for example, are very structurally similar to human hands, and may constitute a [[vestigial structure]]. Other examples include the presence of hip bones in whales and snakes. Such structures may exist with little or no function in a more current organism, yet have a clear function in an ancestral species of the same. Examples of vestigial structures in humans include [[wisdom teeth]], the [[coccyx]] and the [[vermiform appendix]].
[[Phylogenetics]], the study of the ancestry of species, has revealed that structures with similar internal organization may perform divergent functions. [[Vertebrate]] limbs are a common example of such [[homology (biology)|homologous]] structures. The appendages on bat wings, for example, are very structurally similar to human hands, and may constitute a [[vestigial structure]]. Other examples include the presence of hip bones in whales and snakes. Such structures may exist with little or no function in a more current organism, yet have a clear function in an ancestral species of the same. Examples of vestigial structures in humans include [[wisdom teeth]], the [[coccyx]] and the [[vermiform appendix]].


===Molecular evidence===
===Molecular evidence===
Comparison of the DNA sequences allows organisms to be grouped by sequence similarity, and the resulting [[phylogenetics|phylogenetic]] trees are typically congruent with traditional taxonomy, and are often used to strengthen or correct taxonomic classifications. Sequence comparison is considered a measure robust enough to be used to correct erroneous assumptions in the phylogenetic tree in instances where other evidence is scarce.  For example, neutral human DNA sequences are approximately 1.2% divergent (based on substitutions) from those of their nearest genetic relative, the [[chimpanzee]], 1.6% from [[gorilla]]s, and 6.6% from [[baboon]]s.<ref>Two sources: 'Genomic divergences between humans and other hominoids and the effective population size of the common ancestor of humans and chimpanzees'. and 'Quantitative Estimates of Sequence Divergence for Comparative Analyses of Mammalian Genomes' "[http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=11170892] [http://www.genome.org/cgi/content/full/13/5/813]"</ref> Genetic sequence evidence thus allows inference and quantification of genetic relatedness between humans and other apes.<ref>The picture labeled "Human Chromosome 2 and its analogs in the apes" in the article [http://www.gate.net/~rwms/hum_ape_chrom.html Comparison of the Human and Great Ape Chromosomes as Evidence for Common Ancestry] is literally a picture of a link in humans that links two separate chromosomes in the nonhuman apes creating a single chromosome in humans. It is considered a missing link, and the ape-human connection is of particular interest. Also, while the term originally referred to fossil evidence, this too is a trace from the past corresponding to some living beings which when alive were the physical embodiment of this link.</ref><ref>The [[New York Times]] report ''[http://www.nytimes.com/2006/03/07/science/07evolve.html Still Evolving, Human Genes Tell New Story]'', based on ''[http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0040072 A Map of Recent Positive Selection in the Human Genome]'', states the [[International HapMap Project]] is "providing the strongest evidence yet that humans are still evolving" and details some of that evidence.</ref> The sequence of the 16S [[rRNA]] gene, a vital gene encoding a part of the [[ribosome]], was used to find the broad phylogenetic relationships between all extant life.  The analysis, originally done by [[Carl Woese]], resulted in the [[three-domain system]], arguing for two major splits in the early evolution of life.  The first split led to modern [[Bacteria]] and the subsequent split led to modern [[Archaea]] and [[Eukaryote]].   
Comparison of DNA sequences allows organisms to be grouped by sequence similarity, and the resulting [[phylogenetics|phylogenetic]] trees are typically congruent with traditional taxonomy, and are often used to strengthen or correct taxonomic classifications. Sequence comparison is considered a measure robust enough to be used to correct erroneous assumptions in the phylogenetic tree in instances where other evidence is scarce.  For example, neutral human DNA sequences are approximately 1.2% divergent (based on substitutions) from those of their nearest genetic relative, the [[chimpanzee]], 1.6% from [[gorilla]]s, and 6.6% from [[baboon]]s.<ref>Two sources: 'Genomic divergences between humans and other hominoids and the effective population size of the common ancestor of humans and chimpanzees'. and 'Quantitative Estimates of Sequence Divergence for Comparative Analyses of Mammalian Genomes' "[http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=11170892] [http://www.genome.org/cgi/content/full/13/5/813]"</ref> Genetic sequence evidence thus allows inference and quantification of genetic relatedness between humans and other apes.<ref>The picture labeled "Human Chromosome 2 and its analogs in the apes" in the article [http://www.gate.net/~rwms/hum_ape_chrom.html Comparison of the Human and Great Ape Chromosomes as Evidence for Common Ancestry] is literally a picture of a link in humans that links two separate chromosomes in the nonhuman apes creating a single chromosome in humans. It is considered a missing link, and the ape-human connection is of particular interest. Also, while the term originally referred to fossil evidence, this too is a trace from the past corresponding to some living beings which when alive were the physical embodiment of this link.</ref><ref>The [[New York Times]] report ''[http://www.nytimes.com/2006/03/07/science/07evolve.html Still Evolving, Human Genes Tell New Story]'', based on ''[http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0040072 A Map of Recent Positive Selection in the Human Genome]'', states the [[International HapMap Project]] is "providing the strongest evidence yet that humans are still evolving" and details some of that evidence.</ref> The sequence of the 16S [[rRNA]] gene, a vital gene encoding a part of the [[ribosome]], was used to find the broad phylogenetic relationships between all extant life.  The analysis, originally done by [[Carl Woese]], resulted in the [[three-domain system]], arguing for two major splits in the early evolution of life.  The first split led to modern [[Bacteria]] and the subsequent split led to modern [[Archaea]] and [[Eukaryote]].   


The [[Proteome|proteomic]] evidence also supports the universal ancestry of life. Vital [[protein]]s, such as the [[ribosome]], [[DNA polymerase]], and [[RNA polymerase]] are found in the most primitive bacteria to the most complex mammals.  The core part of the protein is conserved across all lineages of life, serving similar functions.  Higher organisms have evolved additional [[protein subunits]], largely affecting the regulation and [[protein-protein interaction]] of the core.  Other overarching similarities between all lineages of extant organisms, such as [[DNA]], [[RNA]], [[amino acids]], and the [[lipid bilayer]], give support to the theory of common descent.  The [[Chirality (chemistry)|chirality]] of [[DNA]], [[RNA]], and [[amino acids]] is conserved across all known life.  As there is no functional advantage to right or left handed molecular [[Chirality (chemistry)|chirality]], the simplest hypothesis is that the choice was made randomly in the early beginnings of life and passed on to all extant life through common descent.
The [[Proteome|proteomic]] evidence also supports the universal ancestry of life. Vital [[protein]]s, such as the [[ribosome]], [[DNA polymerase]], and [[RNA polymerase]] are found in the most primitive bacteria to the most complex mammals.  The core part of the protein is conserved across all lineages of life, serving similar functions.  Higher organisms have evolved additional [[protein subunits]], largely affecting the regulation and [[protein-protein interaction]] of the core.  Other overarching similarities between all lineages of extant organisms, such as [[DNA]], [[RNA]], [[amino acids]], and the [[lipid bilayer]], give support to the theory of common descent.  The [[Chirality (chemistry)|chirality]] of [[DNA]], [[RNA]], and [[amino acids]] is conserved across all known life.  As there is no functional advantage to right or left handed molecular [[Chirality (chemistry)|chirality]], the simplest hypothesis is that the choice was made randomly in the early beginnings of life and passed on to all extant life through common descent.


Molecular evidence also offers a mechanism for large evolutionary leaps and [[macroevolution]].  [[Horizontal gene transfer]], the process in which an organism transfers genetic material (i.e. DNA) to another cell that is not its offspring, allows for large sudden evolutionary leaps in a species by incorporating beneficial genes evolved in another species.  The [[Endosymbiotic theory]] explains the origin of [[mitochondrion|mitochondria]] and [[plastids]] (e.g. [[chloroplast]]s), which are [[organelle]]s of eukaryotic cells, as the incorporation of an ancient [[prokaryote|prokaryotic]] cell into ancient [[eukaryote|eukaryotic]] cell.  Rather than evolving [[eukaryote|eukaryotic]] [[organelle]]s slowly, this theory offers a mechanism for a sudden evolutionary leap by incorporating the genetic material and biochemical composition of a separate species.  This evolutionary mechanism has been observed.  [[Heneta]], a [[protist]], is an extant organism that is undergoing [[Endosymbiotic theory|endosymbiotic]] evolution.<ref>{{cite journal| author = Okamoto N, Inouye I. | year =2005| title = A secondary symbiosis in progress| journal =Science | volume =310 | issue =5746 | pages =287}}</ref><ref>{{cite journal| author = Okamoto N, Inouye I. | year =2006| title = Hatena arenicola gen. et sp. nov., a Katablepharid Undergoing Probable Plastid Acquisition.| journal =Protist |volume = Article in Print}}</ref>     
Molecular evidence also offers a mechanism for large evolutionary leaps and [[macroevolution]].  [[Horizontal gene transfer]], the process in which an organism transfers genetic material (i.e. DNA) to another cell that is not its offspring, allows for large sudden evolutionary leaps in a species by incorporating beneficial genes evolved in another species.  The [[Endosymbiotic theory]] explains the origin of [[mitochondrion|mitochondria]] and [[plastids]] (e.g. [[chloroplast]]s), which are [[organelle]]s of eukaryotic cells, as the incorporation of an ancient [[prokaryote|prokaryotic]] cell into ancient [[eukaryote|eukaryotic]] cell.  Rather than evolving [[eukaryote|eukaryotic]] [[organelle]]s slowly, this theory offers a mechanism for a sudden evolutionary leap by incorporating the genetic material and biochemical composition of a separate species.  This evolutionary mechanism has been observed.  [[Heneta]], a [[protist]], is an extant organism that is undergoing [[Endosymbiotic theory|endosymbiotic]] evolution.<ref>{{cite journal| author = Okamoto N, Inouye I. | year =2005| title = A secondary symbiosis in progress| journal =Science | volume =310 | issue =5746 | pages =287}}</ref><ref>{{cite journal| author = Okamoto N, Inouye I | year =2006| title = Hatena arenicola gen. et sp. nov., a Katablepharid Undergoing Probable Plastid Acquisition.| journal =Protist |volume = }}</ref>     
 
Further evidence for reconstructing ancestral lineages comes from [[junk DNA]] such as [[pseudogene]]s, ''i.e.'', 'dead' genes, which steadily accumulate mutations.<ref>Pseudogene evolution and natural selection for a compact genome. "[http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=10833048]"</ref>
 
Since [[metabolism|metabolic]] processes do not leave fossils, research into the evolution of the basic cellular processes is done largely by comparison of existing organisms. Many lineages diverged when new metabolic processes appeared, and it is theoretically possible to determine when certain metabolic processes appeared by comparing the traits of the descendants of a common ancestor or by detecting their physical manifestations.  As an example, the appearance of oxygen in the earth's atmosphere is linked to the evolution of photosynthesis.
 
===Evidence from studies of complex iteration===
<!--This section uses waaay too many lengthy quotations. Create a daughter article if you want that much level of detail for this section; otherwise, the section should just briefly explain the topic in a few paragraphs, with no more than a couple of brief quotations.-->
"It has taken more than five decades, but the electronic computer is now powerful enough to simulate evolution" assisting [[bioinformatics]] in its attempt to solve biological problems.<ref>[http://www.trnmag.com/Stories/2003/052103/Simulated_evolution_gets_complex_052103.html Simulated evolution gets complex]</ref> [[Computer science]] allows the [[iteration]] of self changing [[complex system]]s to be studied, allowing a mathematically exact understanding of the nature of the processes behind evolution and providing evidence for the hidden causes of known evolutionary events. The evolution of specific cellular mechanisms like [[spliceosome]]s that can turn the cell's genome into a vast workshop of billions of interchangeable parts can be studied for the first time in an exact way.
 
Christoph Adami et al., for example, make this point in ''Evolution of biological complexity'':
 
{{cquote|To make a case for or against a trend in the evolution of complexity in biological evolution, complexity needs to be both rigorously defined and measurable. A recent information-theoretic (but intuitively evident) definition identifies genomic complexity with the amount of information a sequence stores about its environment. We investigate the evolution of genomic complexity in populations of digital organisms and monitor in detail the evolutionary transitions that increase complexity. We show that, because natural selection forces genomes to behave as a natural "Maxwell Demon," within a fixed environment, genomic complexity is forced to increase.<ref>{{cite journal | author=Adami C, Ofria C, Collier TC | title=Evolution of biological complexity | journal=Proc Natl Acad Sci U S A | year=2000 | pages=4463-8 | volume=97 | issue=9 | id=PMID 10781045}}</ref&gt;}}
 
David J. Earl and Michael W. Deem also make this point in ''Evolvability is a selectable trait'':
 
{{cquote|Not only has life evolved, but life has evolved to evolve. That is, correlations within protein structure have evolved, and mechanisms to manipulate these correlations have evolved in tandem. The rates at which the various events within the hierarchy of evolutionary moves occur are not random or arbitrary but are selected by Darwinian evolution. Sensibly, rapid or extreme environmental change leads to selection for greater evolvability. This selection is not forbidden by causality and is strongest on the largest-scale moves within the mutational hierarchy. Many observations within evolutionary biology, heretofore considered evolutionary happenstance or accidents, are explained by selection for evolvability. For example, the vertebrate immune system shows that the variable environment of antigens has provided selective pressure for the use of adaptable codons and low-fidelity polymerases during somatic hypermutation. A similar driving force for biased codon usage as a result of productively high mutation rates is observed in the hemagglutinin protein of [[Influenzavirus A|influenza A]].<ref>{{cite journal | author=Earl DJ, Deem MW | title=Evolvability is a selectable trait | journal=Proc Natl Acad Sci U S A | year=2004 | pages=11531-6 | volume=101 | issue=32 | id=PMID 15289608}}</ref>}}
 
"Computer simulations of the evolution of linear sequences have demonstrated the importance of recombination of blocks of sequence rather than point mutagenesis alone. Repeated cycles of point mutagenesis, recombination, and selection should allow in vitro molecular evolution of complex sequences, such as proteins."<ref>{{cite journal | author=Stemmer WP | title=DNA shuffling by random fragmentation and reassembly: in vitro recombination for molecular evolution | journal=Proc Natl Acad Sci U S A | year=1994 | pages=10747-51 | volume=91 | issue=22 | id=PMID 7938023}}</ref> Evolutionary molecular engineering, also called "directed evolution" or "in vitro molecular evolution", involves the iterated cycle of mutation, multiplication with recombination, and selection of the fittest of individual molecules (proteins, DNA and RNA). The process of natural evolution can be reconstructed, showing possible paths from catalytic cycles based on proteins to ones based on RNA to ones based on DNA.<ref>[http://www.scripps.edu/newsandviews/e_20060327/evo.html scripps.edu]
[http://bio.kaist.ac.kr/~jsrhee/research03.html bio.kaist.ac.kr] [http://www.isgec.org/gecco-2005/free-tutorials.html#ivme free-tutorial] [http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=45099 pubmedcentral.nih.gov]</ref>
 
==Ancestry of organisms==
{{seealso|Common descent}}
[[Image:Huxley - Mans Place in Nature.jpg|left|250px|thumbnail|Morphologic similarities in the [[Hominidae]] family is evidence of common descent.]]
 
In biology, the theory of universal [[common descent]] proposes that all organisms on Earth are descended from a common ancestor or ancestral gene pool.
 
Evidence for common descent is inferred from traits shared between all living organisms. In Darwin's day, the evidence of shared traits was based solely on visible observation of morphologic similarities, such as the fact that all birds, even those which do not fly, have wings. Today, there is strong evidence from genetics that all organisms have a common ancestor. For example, every living cell makes use of [[nucleic acid]]s as its genetic material, and uses the same twenty [[amino acid]]s as the building blocks for [[protein]]s. All organisms use the same [[genetic code]] (with some extremely rare and minor deviations) to [[translation (genetics)|translate]] nucleic acid sequences into proteins. The universality of these traits strongly suggests common ancestry, because the selection of many of these traits seems arbitrary.
 
Information about the early development of life includes input from the fields of geology and [[planetary science]]. These sciences provide information about the history of the Earth and the changes produced by life. However, a great deal of information about the early Earth has been destroyed by geological processes over the course of time.
<br style="clear:both;">
 
===History of life===
<!-- for future reference, heh, here's a ref to stromatolite debate that I took out because it messed up formatting -
"Ancient microfossils from Western Australia are again the subject of heated scientific argument: are they the oldest sign of life on Earth, or just a flaw in the rock?" "[http://www.abc.net.au/science/news/space/SpaceRepublish_497964.htm]" -->
{{main|Timeline of evolution}}
The [[chemical evolution]] from [[Catalyst|self-catalytic chemical reactions]] to [[life]] (see [[Origin of life]]) is not a part of biological evolution, but it is unclear at which point such increasingly complex sets of reactions became what we would consider, today, to be living organisms.
 
[[Image:Stromatolites.jpg|right|thumb|280px|[[Precambrian]] [[stromatolite]]s in the Siyeh Formation, [[Glacier National Park (US)|Glacier National Park]]. In 2002, William Schopf of [[University of California, Los Angeles|UCLA]] published a controversial paper in the journal ''[[Nature (journal)|Nature]]'' arguing that formations such as this possess 3.5 billion year old [[fossil]]ized [[alga]]e microbes. If true, they would be the earliest known life on earth.]]
 
Not much is known about the earliest developments in life. However, all existing organisms share certain traits, including cellular structure and [[genetic code]]. Most scientists interpret this to mean all existing organisms share a common ancestor, which had already developed the most fundamental cellular processes, but there is no [[scientific consensus]] on the relationship of the three domains of life ([[Archaea]], [[Bacterium|Bacteria]], [[Eukaryota]]) or the [[origin of life]]. Attempts to shed light on the earliest history of life generally focus on the behavior of [[macromolecule]]s, particularly [[RNA]], and the behavior of [[complex system]]s.
 
The emergence of oxygenic [[photosynthesis]] (around 3 billion years ago) and the subsequent emergence of an oxygen-rich, non-reducing atmosphere can be traced through the formation of [[Banded iron formation|banded iron]] deposits, and later [[red bed]]s of iron oxides. This was a necessary prerequisite for the development of [[aerobic respiration|aerobic]] [[cellular respiration]], believed to have emerged around 2 billion years ago.
 
In the last billion years, simple multicellular plants and animals began to appear in the oceans. Soon after the emergence of the first animals, the [[Cambrian explosion]] (a period of unrivaled and remarkable, but brief, organismal diversity documented in the fossils found at the [[Burgess Shale]]) saw the creation of all the major body plans, or [[phylum (biology)|phyla]], of modern animals. This event is now believed to have been triggered by the development of the [[Homeobox|Hox genes]]. About 500 million years ago, [[plant]]s and [[fungi]] colonized the land, and were soon followed by [[arthropod]]s and other animals, leading to the development of land [[ecosystem]]s with which we are familiar.
 
The evolutionary process may be exceedingly slow. Fossil evidence indicates that the diversity and complexity of modern life has developed over much of the [[history of Earth|history of the earth]]. [[geology|Geological]] evidence indicates that the Earth is approximately [[Age of the earth|4.6 billion years old]]. Studies on guppies by David Reznick at the University of California, Riverside, however, have shown that the rate of evolution through natural selection can proceed 10 thousand to 10 million times faster than what is indicated in the fossil record.<ref>Evaluation of the Rate of Evolution in Natural Populations of Guppies (Poecilia reticulata) "[http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=9072971&query_hl=2]"</ref>. Such comparative studies however are invariably biased by disparities in the time scales over which evolutionary change is measured in the laboratory, field experiments, and the fossil record.
 
The ancestry of living organisms has traditionally been reconstructed from morphology, but is increasingly supplemented with phylogenetic&mdash;the reconstruction of phylogenies by the comparison of genetic (usually DNA) sequence.<ref>Oklahoma State - [http://opbs.okstate.edu/~melcher/MG/MGW3/MG334.html Horizontal Gene Transfer]: "Sequence comparisons suggest recent horizontal transfer of many [[gene]]s among diverse [[species]] including across the boundaries of [[phylogenetic]] 'domains'. Thus determining the phylogenetic history of a species can not be done conclusively by determining evolutionary trees for single genes."</ref> Biologist Gogarten suggests that "the original metaphor of a tree no longer fits the data from recent genome research", and that therefore "biologists [should] use the metaphor of a mosaic to describe the different histories combined in individual genomes and use [the] metaphor of a net to visualize the rich exchange and cooperative effects of [[horizontal gene transfer|HGT]] among microbes".<ref>[http://www.esalenctr.org/display/confpage.cfm?confid=10&pageid=105&pgtype=1 esalenctr.org]</ref>
 
==Modern synthesis==
{{main|Modern evolutionary synthesis}}
 
Charles Darwin was able to observe variation, infer natural selection and thereby adaptation, but didn't know the basis of heritability.  He couldn't explain ''how'' organisms might change over generations. It also seemed that when two individuals were crossed, their traits must be ''blended'' in the progeny, so that eventually all variation would be lost.
 
The blending problem was solved when the population geneticists [[Ronald Fisher|R.A. Fisher]], [[Sewall Wright]], and [[J. B. S. Haldane]], married Darwinian evolutionary theory to population genetic theory, which was based on [[Gregor Mendel|Mendelian]] [[genetics]] (genes as ''discrete units'' of heredity).
 
The problem of what the mechanisms might be was solved in principle with the identification of DNA as the genetic material by [[Oswald Avery]] and colleagues, and the articulation of the double-helical structure of DNA by [[James Watson]] and [[Francis Crick]] provided a physical basis for the notion that genes were [[genetic code|encoded]] in DNA.
 
===Heredity===
[[Image:DNA123.png|thumb|left|125px|A section of a model of a DNA molecule.]]
 
Gregor Mendel's work provided the first firm basis to the idea that heredity occurred in discrete units. He noticed several traits in peas that occur in only one of two forms (e.g., the peas were either "round" or "wrinkled"), and was able to show that the traits were: heritable (passed from parent to offspring); discrete (i.e., if one parent had round peas and the other wrinkled, the progeny were not intermediate, but either round or wrinkled); and were distributed to progeny in a well-defined and predictable manner ([[Mendelian inheritance]]). His research laid the foundation for the concept of discrete heritable [[Trait (biology)|traits]], known today as [[gene]]s. After Mendel's work was "rediscovered" in 1900, it was discovered that the concepts could have wide applicability, and that most complex traits were polygenetic and not controlled by single unit characters.
 
Later research gave a physical basis to the notion of genes, and eventually identified [[DNA]] as the genetic material, and identified genes as discrete elements within DNA. DNA is not perfectly copied, and rare mistakes ([[mutation]]s) in genes can affect traits that the genes control (e.g., pea shape).
 
A gene can have modifications such as [[DNA methylation]], which do not change the nucleotide sequence of a gene, but do result in the [[epigenetic inheritance]] of a change in the expression of that gene in a trait.
 
Non-DNA based forms of heritable variation exist, such transmission of the secondary structures of [[prion]]s, and [[structural inheritance]] of patterns in the rows of cilia in protozoans such as ''Paramecium''<ref>
BEISSON, J. & SONNEBORN, T. M. (1965). Cytoplasmic inheritance of the organization of the cell cortex of Paramecium aurelia. Proc. natn. Acad Sci. U.S.A. 53, 275-282</ref> and [http://dev.biologists.org/cgi/reprint/105/3/447| ''Tetrahymena'']. Investigations continue into whether these mechanisms allow for the production of specific beneficial heritable variation in response to environmental signals. If this were shown to be the case, then some instances of evolution would lie outside of the typical Darwinian framework, which avoids any connection between environmental signals and the production of heritable variation. However, the processes that produce these variations leave the genetic information intact and are often reversible, and are rather rare.
 
===Variation===
Evolutionary changes are the product of evolutionary forces acting on [[genetic variation]]. In natural populations, there is a certain amount of [[phenotype|phenotypic]] variation (e.g., what makes you appear different from your neighbor). This phenotypic variation is the result of variants in gene sequences among the individuals of a population. There may be one or more functional variants of a gene or locus, and these variants are called [[allele]]s. Most sites in the [[genome]] (''i.e.'', complete DNA sequence) of a species are identical in all individuals in the population; sites with more than one allele are called [[polymorphic]] or segregating sites.
 
All genetic variation begins as a new mutation in a single individual; in subsequent generations the frequency of that variant may fluctuate in the population, becoming more or less prevalent relative to other alleles at the site. This change in allele frequency is the commonly accepted definition of evolution, and all evolutionary forces act by driving allele frequency in one direction or another. Variation disappears when it reaches the point of fixation - when it either reaches a frequency of zero and disappears from the population, or reaches a frequency of one and replaces the ancestral allele entirely.
 
===Mechanisms of evolution===
Evolution consists of two basic types of processes: those that introduce new genetic variation into a population, and those that affect the frequencies of existing variation. Paleontologist [[Stephen J. Gould]] once phrased this succinctly as "''variation proposes and selection disposes.''"<ref>{{cite web|url= http://www.nybooks.com/articles/1151 |title= Darwinian Fundamentalism  |accessdate=2006-08-01 |author= Stephen J. Gould |date=1997-06-12 |publisher=New York Review of Books}}</ref>


These mechanisms of evolution have all been observed in the present and in evidence of their existence in the past. Their study is being used to guide the development of new medicines and other health aids such as the current effort to prevent a [[H5N1]] (i.e. bird flu) pandemic.<ref>The use of evolutionary principles to guide disease diagnosis and drug development with respect to bird flu (i.e. H5N1 virus) is shown [http://www.cdc.gov/ncidod/EID/vol11no10/05-0644.htm here at CDC]. [http://www.nap.edu/books/0309095042/html/123.html#p2000c2099960123001 Here] is the "tree of life" showing the evolution by [[reassortment]] of [[H5N1]] that created the Z genotype in 2002 and [http://www.cdc.gov/ncidod/EID/vol11no10/05-0644-G1.htm here] is evolution by [[antigenic drift]] that created dozens of highly [[pathogenic]] varieties of the Z genotype of avian flu virus [[H5N1]], some of which are increasingly adopted to mammals. Evolution. Right before our eyes.</ref>
Further evidence for reconstructing ancestral lineages comes from so-called [[junk DNA]] such as [[pseudogene]]s, ''i.e.'', 'dead' genes, which steadily accumulate mutations.<ref>Pseudogene evolution and natural selection for a compact genome. "[http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=10833048]"</ref>


====Mutation====
====Mutation====
{{main|Mutation}}
What accounts for genetic variationMutations are permanent, transmissible changes to the [[genetic material]] of a [[cell (biology)|cell]].  These changes can be caused by: "copying errors" in the genetic material during [[cell division]]. (Explain frame shift)
[[Image:dna-split.png|thumb|right|150px|Mutation occurs because of "copy errors" that occur during DNA replication.]]
Exposure to [[Radioactive decay|radiation]], certain chemicals, and some [[virus (biology)|viruses]] can destabilize these genetic molecules and cause biochemical changes that are permanent. When mutations are made in the genes of a cell they ordinarily will be passed on to daughter cells when the cell divides. In multicellular organisms, the mutations will be passed on to progeny if ''germline mutations''that is- if the genes in the [[gametes]] are changed. Mutations that are not affected by natural selection are called [[Neutral theory of molecular evolution|neutral mutations]]. Their frequency in the population is governed by mutation rate, genetic drift and selective pressure on linked alleles. It is understood that most of a species' genome, in the absence of selection, undergoes a steady accumulation of neutral mutations.
Genetic variation arises due to random mutations that occur at a certain rate in the [[genomes]] of all organisms. Mutations are permanent, transmissible changes to the [[genetic material]] (usually [[DNA]] or [[RNA]]) of a [[cell (biology)|cell]], and can be caused by: "copying errors" in the genetic material during [[cell division]]; by exposure to [[Radioactive decay|radiation]], chemicals, or [[virus (biology)|viruses]]. In multicellular organisms, mutations can be subdivided into ''germline mutations'' that occur in the [[gamete]]s and thus can be passed on to progeny, and ''somatic mutations'' that can lead to the malfunction or death of a cell and can cause [[cancer]].
 
Mutations that are not affected by natural selection are called [[Neutral theory of molecular evolution|neutral mutations]]. Their frequency in the population is governed by mutation rate, genetic drift and selective pressure on linked alleles. It is understood that most of a species' genome, in the absence of selection, undergoes a steady accumulation of neutral mutations.
 
Individual [[genes]] can be affected by [[point mutation|point mutations]], also known as [[Single nucleotide polymorphism|SNPs]], in which a single [[base pair]] is altered.  The substitution of a single base pair may or may not affect the function of the gene (see [[mutation]]) while deletions and insertions of a single or several base pairs usually results in a non-functional gene.<ref>Snustad, P. and Simmons, A. 2000. Principles of Genetics, 2nd edition. John Wiley and Sons, Inc. New-York, p.20</ref>


Mobile elements, [[transposons]], make up a major fraction of the genomes of plants and animals and appear to have played a significant role in the evolution of genomes. These mobile insertional elements can jump within a genome and alter existing genes and gene networks to produce evolutionary change and diversity.
Mobile elements, [[transposons]], make up a major fraction of the genomes of plants and animals and appear to have played a significant role in the evolution of genomes. These mobile insertional elements can jump within a genome and alter existing genes and gene networks to produce evolutionary change and diversity.
On the other hand, [[gene duplication]]s, which may occur via a number of mechanisms, are believed to be one major source of raw material for evolving new genes as tens to hundreds of genes are duplicated in animal genomes every million years.<ref>{{cite book| last = Carroll S.B,. Grenier J.K., Weatherbee S.D. | title = From DNA to Diversity: Molecular Genetics and the Evolution of Animal Design.  Second Edition| publisher = Blackwell Publishing| date = 2005| location = Oxford | id = ISBN 1-4051-1950-0 }}</ref> Most genes belong to larger "families" of genes derived from a common ancestral gene (two genes from a species that are in the same family are dubbed "[[paralog]]s"). Another mechanism causing gene duplication is intergenic recombination, particularly '[http://www.pnas.org/cgi/content/full/94/15/7698| exon shuffling]', ''i.e.'', an abberant recombination that joins the 'upstream' part of one gene with the 'downstream' part of another.  [[Polyploidy|Genome duplications]] and chromosome duplications also appear to have served a significant role in evolution.  Genome duplication has been the driving force in the [[Teleostei]] genome evolution, where up to four genome duplications are thought to have happened, resulting in species with more than 250 chromosomes.
Large [[chromosomal]] rearrangements do not necessarily change gene function, but do generally result in reproductive isolation, and, by definition, [[speciation]] ([[species]] (in sexual organisms) are usually defined by the ability to interbreed).  An example of this mechanism is the fusion of two chromosomes in the [[homo genus]] that produced human chromosome 2; this fusion did not occur in the [[chimp]] [[Lineage (evolution)|lineage]], resulting in two separate chromosomes in extant chimps.


====Selection and adaptation====
====Selection and adaptation====
{{main|Natural selection|Adaptation}}
{{main|Natural selection|Adaptation}}
[[Image:Peacock.displaying.better.800pix.jpg|thumb|right|250px|A [[peacock]]'s tail is the canonical example of [[sexual selection]]]]


Natural selection comes from differences in survival and reproduction <!-- The environment is just one of the two major sources (ecological selection), the other is partners (sexual selection)-->. Differential mortality is the survival rate of individuals to their reproductive age. Differential fertility is the total genetic contribution to the next generation. Note that, whereas mutations and genetic drift are random, natural selection is not, as it preferentially selects for different mutations based on differential fitnesses. For example, rolling dice is random, but always picking the higher number on two rolled dice is not random. The central role of natural selection in evolutionary theory has given rise to a strong connection between that field and the study of [[ecology]].


Natural selection can be subdivided into two categories:
Natural selection can be subdivided into two categories:
Line 188: Line 110:
** [[Genetic hitchhiking]] occurs when a beneficial [[allele]] is selected for, and [[Genetic linkage|linked]] [[allele]]s, which can be neutral or beneficial, are pushed towards fixation along with the beneficial [[allele]].
** [[Genetic hitchhiking]] occurs when a beneficial [[allele]] is selected for, and [[Genetic linkage|linked]] [[allele]]s, which can be neutral or beneficial, are pushed towards fixation along with the beneficial [[allele]].


Through the process of natural selection, organisms become better adapted to their environments. [[Adaptation]] is any evolutionary process that increases the [[fitness (biology)|fitness]] of the individual, or sometimes the trait that confers increased fitness, e.g. a stronger prehensile tail or greater visual acuity. Note that adaptation is context-sensitive; a trait that increases fitness in one environment may decrease it in another.
Evolution does not act in a linear direction towards a pre-defined "goal" &mdash; it only responds to various types of adaptionary changes. The belief in a [[teleology|telelogical]] evolution of this sort is known as [[orthogenesis]], and is not supported by the scientific understanding of evolution. One example of this misconception is the erroneous belief humans will evolve [[polydactyly|more fingers]] in the future on account of their increased use of machines such as [[computer]]s. In reality, this would only occur if more fingers offered a significantly higher rate of reproductive success than those not having them, which seems very unlikely at the current time.


Most biologists believe that adaptation occurs through the accumulation of many mutations of small effect. However, [[macromutation]] is an alternative process for adaptation that involves a single, very large scale mutation.
Most biologists believe that adaptation occurs through the accumulation of many mutations of small effect. However, [[macromutation]] is an alternative process for adaptation that involves a single, very large scale mutation.


====Recombination====
====Recombination====
{{Main|Genetic recombination}}
In asexual organisms, variants in genes on the same chromosome will always be inherited together - they are '''linked''', by virtue of being on the same DNA molecule. However, [[sex|sexual]] organisms, in the production of gametes, shuffle linked alleles on homologous chromosomes inherited from the parents via [[meiosis|meiotic]] [[recombination]]. This shuffling allows independent [[Mendelian inheritance#Mendel.27s law of segregation|assortment]] of alleles (mutations) in genes to be propagated in the population independently. This allows bad mutations to be purged and beneficial mutations to be retained more efficiently than in asexual populations.
In asexual organisms, variants in genes on the same chromosome will always be inherited together - they are '''linked''', by virtue of being on the same DNA molecule. However, [[sex|sexual]] organisms, in the production of gametes, shuffle linked alleles on homologous chromosomes inherited from the parents via [[meiosis|meiotic]] [[recombination]]. This shuffling allows independent [[Mendelian inheritance#Mendel.27s law of segregation|assortment]] of alleles (mutations) in genes to be propagated in the population independently. This allows bad mutations to be purged and beneficial mutations to be retained more efficiently than in asexual populations.


However, the meitoic recombination rate is not very high - on the order of one crossover (recombination event between homomolgous chromosomes) per chromosome arm per generation. Therefore, linked alleles are not perfectly shuffled away from each other, but tend to be inherited together. This tendency may be measured by comparing the co-occurrence of two alleles, usually quantified as [[linkage disequilibrium]] (LD). A set of alleles that are often co-propagated is called a [[haplotype]]. Strong haplotype blocks can be a product of strong positive selection.
However, the meitoic recombination rate is not very high - on the order of one crossover (recombination event between homomolgous chromosomes) per chromosome arm per generation. Therefore, linked alleles are not perfectly shuffled away from each other, but tend to be inherited together. This tendency may be measured by comparing the co-occurrence of two alleles, usually quantified as [[linkage disequilibrium]] (LD). A set of alleles that are often co-propagated is called a [[haplotype]]. Strong haplotype blocks can be a product of strong positive selection.


Recombination is mildly mutagenic, which is one of the proposed reasons why it occurs with limited frequency. Recombination also breaks up gene combinations that have been successful in previous generations, and hence should be opposed by selection. However, recombination could be favoured by negative frequency-dependent selection (this is when rare variants increase in frequency) because it leads to more individuals with new and rare gene combinations being produced.
====Gene flow and population structure====
 
When alleles cannot be separated by recombination (for example in mammalian [[Y chromosome]]s), we see a reduction in [[effective population size]], known as the [[Hill Robertson effect]], and the successive establishment of bad mutations, known as [[Muller's ratchet]].
 
====Gene flow and Population structure====
:''Main article: [[Population genetics]]''
 
[[Image:Evolution_evi_mig.png|350px|thumb|right|Map of the world showing distribution of [[camelid]]s. Solid black lines indicate possible migration routes.]]
 
[[Gene flow]] (also called ''gene admixture'' or simply ''migration'') is the exchange of genetic variation between populations, when geography and culture are not obstacles. [[Ernst Mayr]] thought that gene flow is likely to be homogenising, and therefore counteract selective adaptation. Where there are obstacles to gene flow, the situation is termed [[reproductive isolation]] and is considered to be necessary for [[speciation]].
[[Gene flow]] (also called ''gene admixture'' or simply ''migration'') is the exchange of genetic variation between populations, when geography and culture are not obstacles. [[Ernst Mayr]] thought that gene flow is likely to be homogenising, and therefore counteract selective adaptation. Where there are obstacles to gene flow, the situation is termed [[reproductive isolation]] and is considered to be necessary for [[speciation]].


Line 217: Line 126:


====Drift====
====Drift====
{{main|Genetic drift}}
[[Genetic drift]] describes changes in allele frequency from one generation to the next due to [[variance|sampling variance]]. The frequency of an allele in the offspring generation will vary according to a probability distribution of the frequency of the allele in the parent generation. Thus, over time even in the absence of selection upon the alleles, allele frequencies will tend to "drift" upward or downward, eventually becoming "fixed" - that is, going to 0% or 100% frequency. Thus, fluctuations in allele frequency between successive generations may result in some alleles disappearing from the population due to chance alone. Two separate populations that begin with the same allele frequencies therefore might drift  apart by random fluctuation into two divergent populations with different allele sets (for example, alleles present in one population could be absent in the other, or ''vice versa'').
 
Genetic drift describes changes in allele frequency from one generation to the next due to [[variance|sampling variance]]. The frequency of an allele in the offspring generation will vary according to a probability distribution of the frequency of the allele in the parent generation. Thus, over time even in the absence of selection upon the alleles, allele frequencies will tend to "drift" upward or downward, eventually becoming "fixed" - that is, going to 0% or 100% frequency. Thus, fluctuations in allele frequency between successive generations may result in some alleles disappearing from the population due to chance alone. Two separate populations that begin with the same allele frequencies therefore might drift  apart by random fluctuation into two divergent populations with different allele sets (for example, alleles present in one population could be absent in the other, or ''vice versa'').


The impact of genetic drift depends strongly on the size of the population (generally abbreviated as N): drift is important in small mating populations (see [[Founder effect]] and [[Population bottleneck]]), where chance fluctuations from generation to generation can be large. The relative importance of natural selection and genetic drift in determining the fate of new mutations also depends on the population size and the strength of selection: when N times s (population size times strength of selection) is small, genetic drift predominates. When N times s is large, selection predominates. Thus, natural selection is predominant in large populations, or equivalently, genetic drift is stronger in small populations. Finally, the time for an allele to become fixed in the population by genetic drift (that is, for all individuals in the population to carry that allele) depends on population size, with smaller populations requiring a shorter time to fixation.
The impact of genetic drift depends strongly on the size of the population (generally abbreviated as N): drift is important in small mating populations (see [[Founder effect]] and [[Population bottleneck]]), where chance fluctuations from generation to generation can be large. The relative importance of natural selection and genetic drift in determining the fate of new mutations also depends on the population size and the strength of selection: when N times s (population size times strength of selection) is small, genetic drift predominates. When N times s is large, selection predominates. Thus, natural selection is predominant in large populations, or equivalently, genetic drift is stronger in small populations. Finally, the time for an allele to become fixed in the population by genetic drift (that is, for all individuals in the population to carry that allele) depends on population size, with smaller populations requiring a shorter time to fixation.
====Horizontal gene transfer====
{{main|Horizontal gene transfer}}
[[Horizontal gene transfer]] (HGT) (or Lateral gene transfers) is any process in which an organism transfers [[genetic material]] (i.e. [[DNA]]) to another organism that is not its offspring.  This mechanism allows for the transfer of genetic material between unrelated organisms of the same [[species]] or of different species.
Many mechanisms for [[horizontal gene transfer]] have been observed, such as [[antigenic shift]], [[reassortment]], and [[hybrid]]ization. Viruses can transfer genes between species via [[Transduction (genetics)|transduction]],<ref>[http://66.102.7.104/search?q=cache:tpICVNWaTbgJ:non.fiction.org/lj/community/ref_courses/3484/enmicro.pdf+sex+evolution+%22Horizontal+gene+transfer%22+-human+Conjugation+RNA+DNA&hl=en enmicro.pdf]</ref>. Bacteria can incorporate genes from other dead bacteria or [[plasmids]] via [[Transformation (genetics)|transformation]], exchange genes with living bacteria via [[Bacterial conjugation|conjugation]], and can have [[plasmid]]s "set up residence separate from the host's genome".<ref>[http://www2.nau.edu/~bah/BIO471/Reader/Pennisi_2003.pdf Pennisi_2003.pdf]</ref> 
HGT has been shown to result in the spread of [[antibiotic resistance]] across [[bacterial]] populations.<ref>{{cite journal| last = Dzidic S, Bedekovic V.| title = Horizontal gene transfer-emerging multidrug resistance in hospital bacteria | journal = Acta pharmacologica Sinica| volume = 24| issue = 6 | pages = 519-526 | date = 2003}}</ref>  Furthermore, findings indicate that HGT has been a major mechanism for [[prokaryotic]] and [[eukaryotic]] evolution.<ref>{{cite journal| last = Andersson JO| title = Lateral gene transfer in eukaryotes | journal = Cellular and molecular life sciences| volume = 62| issue = 11 | pages = 1182-1197 | date = 2005}}</ref><ref>{{cite journal| last = Katz LA| title = Lateral gene transfers and the evolution of eukaryotes: theories and data | journal = International journal of systematic and evolutionary microbiology| volume = 52| issue = 5 | pages = 1893-1900 | date = 2002}}</ref>
HGT complicates the inference of the [[phylogeny]] of life, as the original metaphor of a tree of life no longer fits.  Rather, since genetic information is passed to other organisms and other species in addition to being passed from parent to offspring, ''"biologists [should] use the metaphor of a mosaic to describe the different histories combined in individual genomes and use [the] metaphor of a net to visualize the rich exchange and cooperative effects of HGT among microbes."''<ref>[http://www.esalenctr.org/display/confpage.cfm?confid=10&pageid=105&pgtype=1 Evolutionary Theory] by Peter Gogarten, Ph.D.</ref>


===Speciation and extinction===
===Speciation and extinction===
[[Image:Allosaurus1.jpg|right|thumb|200px|An [[Allosaurus]] skeleton.]]


[[Speciation]] is the process by which new biological species arise. This may take place by various mechanisms. [[Allopatric speciation]] occurs in populations that become isolated geographically, such as by [[habitat fragmentation]] or migration.<ref>{{cite journal| author =Hoskin ''et al'' | year =Oct 2005| title = Reinforcement drives rapid allopatric speciation| journal =Nature | volume =437 |  pages =1353-1356}}</ref>. [[Sympatric speciation]] occurs when new species emerge in the same geographic area<ref>{{cite journal| author =Savolainen ''et al'' | year =May 2006| title = Sympatric speciation in palms on an oceanic island| journal =Nature | volume =441 |  pages =210-213}}</ref><ref>{{cite journal| author =Barluenga ''et al'' | year =February 2006| title = Sympatric speciation in Nicaraguan crater lake cichlid fish| journal =Nature | volume =439 | pages =719-723}}</ref> [[Ernst Mayr]]'s [[peripatric speciation]] is a type of speciation that exists in between the extremes of allopatry and sympatry. Peripatric speciation is a critical underpinning of the theory of [[punctuated equilibrium]]. An example of rapid sympatric speciation can be eloquently represented in the [[triangle of U]]; where new species of ''Brassica sp.'' have been made by the fusing of separate genomes from related plants.  
[[Speciation]] is the process by which new biological species arise. This may take place by various mechanisms. [[Allopatric speciation]] occurs in populations that become isolated geographically, such as by [[habitat fragmentation]] or migration.<ref>{{cite journal| author =Hoskin ''et al'' | year =Oct 2005| title = Reinforcement drives rapid allopatric speciation| journal =Nature | volume =437 |  pages =1353-1356}}</ref>. [[Sympatric speciation]] occurs when new species emerge in the same geographic area<ref>{{cite journal| author =Savolainen ''et al'' | year =May 2006| title = Sympatric speciation in palms on an oceanic island| journal =Nature | volume =441 |  pages =210-213}}</ref><ref>{{cite journal| author =Barluenga ''et al'' | year =February 2006| title = Sympatric speciation in Nicaraguan crater lake cichlid fish| journal =Nature | volume =439 | pages =719-723}}</ref> [[Ernst Mayr]]'s [[peripatric speciation]] is a type of speciation that exists in between the extremes of allopatry and sympatry. Peripatric speciation is a critical underpinning of the theory of [[punctuated equilibrium]]. An example of rapid sympatric speciation can be eloquently represented in the [[triangle of U]]; where new species of ''Brassica sp.'' have been made by the fusing of separate genomes from related plants.  
Line 241: Line 137:
{{-}}
{{-}}


==Current Research==
==Current research==
{{seealso|Ancestral Reconstruction|Human Genome Project|Bioinformatics|Evo-devo}}
Evolution is still an active field of research in the scientific community.  Improvements in sequencing methods have resulted in a large increase of sequenced genomes, allowing for the testing and refining of the theory of evolution with respect to whole genome data.  Advances in computational hardware and software have allowed for the testing and extrapolation of increasingly advanced evolutionary models.  Discoveries in biotechnology have produced methods for the ‘’de novo’’ synthesis of proteins and, potentially, entire genomes, driving evolutionary studies at the molecular level.
Evolution is still an active field of research in the scientific community.  Improvements in sequencing methods have resulted in a large increase of sequenced genomes, allowing for the testing and refining of the theory of evolution with respect to whole genome data.  Advances in computational hardware and software have allowed for the testing and extrapolation of increasingly advanced evolutionary models.  Discoveries in biotechnology have produced methods for the ‘’de novo’’ synthesis of proteins and, potentially, entire genomes, driving evolutionary studies at the molecular level.


===Micro RNA's===
==Common misunderstandings about evolutionary theory==
Small RNA's or micro RNA's ([[miRNA]]) appear highly significant in regulation of gene expression during development. They support the [[RNA world hypothesis]] and serve a role in the evolution of plants and animals.<ref>{{cite journal | author = Sempere LF, Cole CN, McPeek MA, Peterson KJ.| title = The phylogenetic distribution of metazoan microRNAs: insights into evolutionary complexity and constraint..| journal = J Exp Zoolog B Mol Dev Evol | volume = | issue = Jul 12 | pages =  | year =2006 | id = PMID 16838302}}</ref><ref>{{cite journal | author = Massirer KB, Pasquinelli AE.| title = The evolving role of microRNAs in animal gene expression.journal = Bioessays.  | volume = 28 | issue =5 | pages = 449-52 | year =2006 | id = PMID 16615087}}</ref><ref>{{cite journal | author = Zhang B, Pan X, Cannon CH, Cobb GP, Anderson TA.| title = Conservation and divergence of plant microRNA genes.| journal = Plant J. | volume = 46 | issue =2 | pages = 243-59 | year =2006 | id = PMID 16623887}}</ref> Micro RNA's contribution to evolution is considered an [[epigenetic]] mechanism in [[evolutionary developmental biology]]. Micro RNA's appear to constitute 1% of the human genome. Scientist are designing silencing interference micro RNA's in the hopes of shutting down genes involved in cancer, diseases, and the contribution of genes in developmental biology.
===Progressiveness, complexity, and evolution===
{|align="right" cellpadding="10" style="background:lightgray; width:25%; border: 1px solid #aaa; margin:20px; font-size: 92%; font-family: Gill Sans MT;"


==Misunderstandings about modern evolutionary biology==
| Evolutionary processes and, in general, scientific explanations of the world are often in contrast with the immediate and simple explanations that our brain gives of reality (e.g. the sun seems to turn around the earth, the earth seems to be flat), and are influenced by what Francis Bacon called "idola"[<ref name=hall>Hall MP. [http://www.sirbacon.org/links/4idols.htm The Four Idols of Francis Bacon: The New Instrument of Knowledge].
 
*<font face="Gill Sans MT">"In the Novum Organum (the new instrumentality for the acquisition of knowledge) Francis Bacon classified the intellectual fallacies of his time under four headings which he called idols. He distinguished them as idols of the Tribe, idols of the Cave, idols of the Marketplace and idols of the Theater…An idol is an image, in this case held in the mind, which receives veneration but is without substance in itself. Bacon did not regard idols as symbols, but rather as fixations."</font></ref>] (false notions or tendencies which distort the truth [<ref>Fantini F. (2005) Didattica dell'evoluzione. In Evoluzione tra ricerca e didattica, XIV – Special number Edited by: Associazione Nazionale Insegnanti di Scienze Naturali. Agnano Pisano: Stamperia Editoriale Pisana; 2005:203-209.</ref>]).<ref name=guidetti>Guidetti R, Baraldi L, Calzolai C, Pini L, Veronesi P, Pederzoli A. (2007) [http://www.biomedcentral.com/1471-2148/7/S2/S Fantastic animals as an experimental model to teach animal adaptation]. ''BMC Evolutionary Biology'' 7(Suppl 2):S13 doi: 10.1186/1471-2148-7-S2-S13.</ref>
Although the modern synthesis is a central theory in science, many misunderstandings about specific facets of the modern synthesis are prevalent among the general population. These misunderstandings have had a negative impact on the acceptance of the modern synthesis,<ref>[http://news.bbc.co.uk/2/hi/education/817000.stm BBC Report on Biology Education in North America] "''In a study of 1,200 college freshmen, Professor Alters found that 45% of those who doubted the theory of evolution had specific misunderstandings about some of the science that has been used to support it.''"</ref><ref>{{cite journal| author =Constance Holden| year =1998| title = SCIENCE EDUCATION: Academy Rallies Teachers on Evolution| journal =Science | volume =280 | issue =5361 | pages =194}}</ref> most notably in the [[United States]].<ref>{{cite journal| author =Miller JD,Scott EC, Okamoto S.| year =2006| title = Science communication. Public acceptance of evolution.| journal =Science | volume =313 | issue =5788 | pages =765-766}}</ref> Some of the most common misunderstandings are examined in this section.  
|}
 
One of the most common misunderstandings of evolution is that one species can be "more highly evolved" than another, that evolution is necessarily progressive and/or leads to greater "complexity", or that its converse is "[[devolution (fallacy)|devolution]]".<ref>[http://www.talkorigins.org/indexcc/CB/CB932.html talkorigins Claim CB932: Evolution of degenerate forms]</ref> Evolution provides no assurance that later generations are more intelligent or complex than earlier generations. The claim that evolution results in progress is not part of modern evolutionary theory; it derives from earlier belief systems which were held around the time Darwin devised his theory of evolution.
===Distinctions between theory and fact===
:''Further information: [[Theory#Science|Scientific Theory]]
: ''See also'': [[Creationism versus evolution#Theory vs. fact|Theory vs. Fact]]
[[Stephen Jay Gould]] explained that "evolution is a theory. It is also a fact. And facts and theories are different things, not rungs in a hierarchy of increasing certainty. Facts are the world's data. Theories are structures of ideas that explain and interpret facts. Facts do not go away when scientists debate rival theories to explain them. Einstein's theory of gravitation replaced Newton's, but apples did not suspend themselves in mid-air, pending the outcomeAnd humans evolved from ape-like ancestors whether they did so by Darwin's proposed mechanism or by some other yet to be discovered."<ref>[http://www.stephenjaygould.org/library/gould_fact-and-theory.html Stephen Jay Gould, "Evolution as Fact and Theory" 1994]</ref>
 
The modern synthesis, like its Mendelian and Darwinian antecedents, is a ''scientific theory.''  A theory is an attempt to identify and describe relationships between phenomena or things, and generates [[falsifiability|falsifiable]] predictions which can be tested through [[scientific method|controlled experiments]] and [[empiricism|empirical observation]]. Speculative or conjectural explanations tend to be called [[hypothesis|hypotheses]], and well tested explanations, ''theories''. [[fact#"Fact" in science|''Fact'']] tends to mean a ''datum'', an ''observation'', ''i.e.'', a fact is obtained by a fairly direct observation. However, a fact does not mean absolute certainty; in science, fact can only mean "confirmed to such a degree that it would be perverse to withhold provisional assent."<ref>[http://www.stephenjaygould.org/library/gould_fact-and-theory.html Stephen Jay Gould, "Evolution as Fact and Theory" 1994]</ref> A ''theory'' is obtained by inference from a body of ''facts''. [[Fact]] and [[theory]] denote the [[epistemology|epistemological status]] of knowledge; how the knowledge was obtained, what sort of knowledge it is
 
In this [[science|scientific]] sense, "facts" are what theories attempt to explain. So, for scientists "theory" and "fact" do not stand in opposition, but rather exist in a reciprocal relationship; for example, it is a "fact" that an apple will fall to the ground if it becomes dislodged from a branch and the "theory" which explains this is the current theory of [[gravitation]]. In the same way, heritable variation, natural selection, and response to selection (''e.g.'' in domesticated plants and animals) are "facts", and the generalization or extrapolation beyond these phenomena, and the explanation for them, is the "theory of evolution".<ref>[http://www.talkorigins.org/faqs/evolution-fact.html Evolution is a Fact and a Theory]</ref>


===Evolution and devolution===
Evolution has involved "progression" towards more complexity, since all of the earliest lifeforms were extremely simple and there was nowhere to go but up. However, there is no guarantee that any particular organism existing today will become more intelligent, more complex, bigger, or stronger in the future. In fact, natural selection will only favor this kind of "progression" if it has a variant to act upon that increases chance of survival, i.e. the ability to live long enough to raise offspring to [[sexual maturity]]. The same mechanism can actually favor lower intelligence, lower complexity, and so on if those traits become a selective advantage in the organism's environment.
One of the most common misunderstandings of evolution is that one species can be "more highly evolved" than another, that evolution is necessarily progressive and/or leads to greater "complexity", or that its converse is "[[devolution (fallacy)|devolution]]".<ref>[http://www.talkorigins.org/indexcc/CB/CB932.html talkorigins Claim CB932: Evolution of degenerate forms]</ref> Evolution provides no assurance that later generations are more intelligent or complex than earlier generations. The claim that evolution results in progress is not part of modern evolutionary theory; it derives from earlier belief systems which were held around the time Darwin devised his theory of evolution.


In many cases evolution does involve "progression" towards more complexity, since the earliest lifeforms were extremely simple compared to many of the species existing today, and there was nowhere to go but up. However, there is no guarantee that any particular organism existing today will become more intelligent, more complex, bigger, or stronger in the future. In fact, natural selection will only favor this kind of "progression" if it increases chance of survival, i.e. the ability to live long enough to raise offspring to [[sexual maturity]]. The same mechanism can actually favor lower intelligence, lower complexity, and so on if those traits become a selective advantage in the organism's environment. One way of understanding the apparent "progression" of lifeforms over time is to remember that the earliest life began as maximally simple forms. Evolution caused life to become more complex, since becoming simpler wasn't advantageous. Once individual lineages have attained sufficient complexity, however, simplifications ([[Specialization (functional)|specialization]]) are as likely as increased complexity. This can be seen in many parasite species, for example, which have evolved simpler forms from more complex ancestors.<ref>[http://www.sciam.com/askexpert_question.cfm?articleID=00071863-683B-1C72-9EB7809EC588F2D7 Scientific American; Biology: Is the human race evolving or devolving?]</ref>
One way of understanding the apparent "progression" of lifeforms over time is to remember that the earliest life began as components of cells. Evolution caused life to become more complex, since becoming simpler wasn't advantageous. Once individual lineages have attained sufficient complexity, simplifications ([[Specialization (functional)|specialization]]) become more likely.


===Speciation===
===Speciation===
{{main|Speciation}}
[[Image:Darwin's finches.jpeg|frame|right|The existence of several different, but related, finches on the [[Galápagos Islands]] is evidence of the occurrence of speciation.]]
It is sometimes claimed that [[speciation]] &ndash; the origin of new species &ndash; has never been directly observed, and thus evolution cannot be called sound science. This is a misunderstanding of both science and evolution. First, scientific discovery does not occur solely through [[Reproducibility|reproducible]] [[experiment]]s; the principle of [[Uniformitarianism (science)|uniformitarianism]] allows natural scientists to infer causes through their empirical effects. Moreover, since the publication of ''On the Origin of Species'' scientists have confirmed Darwin's hypothesis by data gathered from sources that did not exist in his day, such as [[DNA]] similarity among species and new [[Fossil record|fossil]] discoveries. Finally, speciation has actually been directly observed.<ref>{{cite web|url=http://www.talkorigins.org/faqs/faq-speciation.html#part5|title=Observed Instances of Speciation|publisher=Talk Origins Archive|first=Joseph|last=Boxhorn}}</ref> (See the [[Evidence of evolution#Hawthorn fly|hawthorn fly]] example.)
It is sometimes claimed that [[speciation]] &ndash; the origin of new species &ndash; has never been directly observed, and thus evolution cannot be called sound science. This is a misunderstanding of both science and evolution. First, scientific discovery does not occur solely through [[Reproducibility|reproducible]] [[experiment]]s; the principle of [[Uniformitarianism (science)|uniformitarianism]] allows natural scientists to infer causes through their empirical effects. Moreover, since the publication of ''On the Origin of Species'' scientists have confirmed Darwin's hypothesis by data gathered from sources that did not exist in his day, such as [[DNA]] similarity among species and new [[Fossil record|fossil]] discoveries. Finally, speciation has actually been directly observed.<ref>{{cite web|url=http://www.talkorigins.org/faqs/faq-speciation.html#part5|title=Observed Instances of Speciation|publisher=Talk Origins Archive|first=Joseph|last=Boxhorn}}</ref> (See the [[Evidence of evolution#Hawthorn fly|hawthorn fly]] example.)


A variation of this assertion that [[microevolution]] has been observed and [[macroevolution]] has not been observed is subject to the same counterarguments. While there is debate over the details of how evolution happened, it is generally accepted that macroevolution uses the same mechanisms of change as those already observed in microevolution.
A variation of this assertion that [[microevolution]] has been observed and [[macroevolution]] has not been observed is subject to the same counterarguments. While there is debate over the details of how evolution happened, it is generally accepted that macroevolution uses the same mechanisms of change as those already observed in microevolution.


===Self-organization and entropy===
===Purposiveness and evoluton===
{{main|Self-organization}}


It is claimed that evolution, by increasing complexity without supernatural intervention, violates the [[second law of thermodynamics]]. This [[Physical law|law]] posits that in an idealised [[isolated system]], [[entropy]] will tend to increase or stay the same. Entropy is a measure of the dispersal of energy in a physical system so that it is not available to do mechanical work.<ref name=crutch>[http://www.entropysite.com/cracked_crutch.html Disorder — A Cracked Crutch For Supporting Entropy Discussions]</ref> The claim ignores the fact that biological systems are not [[isolated system]]s.  Simple calculations<ref>[http://physics.gmu.edu/~roerter/EvolutionEntropy.htm Does Life On Earth Violate the Second Law of Thermodynamics?]</ref> show that the Sun-Earth-space system does ''not'' violate the second law because the massive increase in entropy due to the Sun and Earth radiating into space dwarfs the small decrease in entropy caused by the evolution of life. In [[statistical thermodynamics]] entropy has been envisioned as a measure of the statistical "disorder" at a [[Microstate (statistical mechanics)|microstate level]], leading to the mistaken idea that entropy implies increasing chaos.<ref name=crutch/>
Peter Bowler, eminent historian of science, in his 3rd edition of his widely cited ''Evolution: History of an Idea'', writes:


In fact, the flow of matter and energy through [[open system (system theory)|open system]]s allows [[self-organization]] enabling an increase in complexity without guidance or management. Examples include mineral crystals and snowflakes. Life inherently involves open systems, not isolated systems, as all organisms exchange energy and matter with their environment, and similarly the Earth receives energy from the Sun and emits energy back into space.
{|align="center" style="width:90%;font-size:98%;"
|
<font face="Gill Sans MT">The antagonism of the creationists should not blind us to the fact that science and religion have sometimes been able to work in harmony. The history of evolutionism reveals many attempts to see the development of life on earth as the unfolding of a divine plan. There have certainly been some scientists who would adopt a more militant posture, arguing that humanity simply has to come to terms with the unpleasant fact that it is the product of a purposeless sequence of natural events.</font><ref name=bowler2003/>
|}


===Information===
Yes, we humans may have resulted from “a purposeless sequence of natural events”, but the fact of that happening to us I do not find “unpleasant”. The purposelessness of the sequence of natural events in evolution, even considering biological evolution only, does not necessarily exclude, in a lineage of entities, that new kinds of evolvable entities might emerge that could impose purpose on evolutionary change. Indeed, human culture emerged, in part from new entities called ‘memes’ or ‘culturgens’, whose evolvable complex associations create knowledge and ideas. The ''eugenics movement'' early exemplified the potentiality for the lineage culminating in ''Homo sapiens'' to ''purposefully'' order the sequence of natural events.
Some assert that evolution cannot create information, or that information can only be created by an intelligence. [[Physical information]] exists regardless of the presence of an intelligence, and evolution allows for new information whenever a novel mutation or [[gene]] duplication occurs and is kept. It does not need to be beneficial or visually apparent to be "information." However, even if those were requirements they would be satisfied with the appearance of [[nylon]]-eating [[bacteria]],<ref>{{cite web|url=http://www.nmsr.org/nylon.htm|title=Evolution and Information: The Nylon Bug|publisher=New Mexicans for Science and Reason}}</ref> which required new [[enzyme]]s to efficiently digest a material that never existed until the modern age.<ref>It wasn't a highly competent design because the bacteria weren't extracting a lot of energy from the process, just enough to get by. And it was based on a simply frame shift reading of a gene that had other uses. But with a simple frame shift of a gene that was already there, it could now "eat" nylon. Future mutations, perhaps point mutations inside that gene, could conceivably heighten the energy gain of the nylon decomp process, and allow the bacteria to truly feast and reproduce faster and more plentifully on just nylon, thus leading perhaps in time to an irreducibly complex arrangement between bacteria who live solely on nylon and a man-made fiber produced only by man. [http://www.edwardtbabinski.us/evolution/darwin_design.html Darwinism or Directed Mutations?]</ref>
 
Japanese researchers demonstrated that nylon degrading ability can be obtained ''[[de novo]]'' in laboratory cultures of ''Pseudomonas aeruginosa'' strain POA, which initially had no enzymes capable of degrading nylon oligomers. This indicates that the ability of bacteria to digest nylon can evolve if proper artificial selection is applied.<ref>{{cite journal|author=Prijambada I.D. ''et al'' | year =1995 | title = Emergence of nylon oligomer degradation enzymes in Pseudomonas aeruginosa PAO through experimental evolution| journal =Applied and Environmental Microbiology | volume = 61| issue =5 | pages = 2020–2022}}</ref> Recently, the same group solved the high resolution [[X-ray crystallography|X-ray crystal structure]] of the newly evolved nylon-digesting [[enzyme]].<ref>{{cite journal|author=Negoro S ''et al'' | year =2005 | title =X-ray crystallographic analysis of 6-aminohexanoate-dimer hydrolase: molecular basis for the birth of a nylon oligomer-degrading enzyme| journal =The Journal of Biological Chemistry | volume = 280| issue =47 | pages = 39644-39652}}</ref> Using the structural results, the authors propose "that the amino acid replacements in the catalytic cleft of a preexisting esterase with the beta-lactamase fold resulted in the evolution of the" nylon-digesting [[enzyme]]. This hypothesis still needs to be confirmed by detailed mutagenesis studies.


==Social and religious controversies==
==Social and religious controversies==
{{main|Social effect of evolutionary theory|Creation-evolution controversy}}
[[Image:Darwin ape.jpg|left|150px|thumb|A satirical 1871 image of [[Charles Darwin]] as a quadrupedal [[ape]] reflects part of the social controversy over whether humans and other apes share a common lineage.]]
Starting with the publication of ''[[The Origin of Species]]'' in 1859, the modern science of evolution has been a source of nearly constant controversy. In general, controversy has centered on the philosophical, [[cosmology|cosmological]], social, and religious implications of evolution, not on the science of evolution itself. The proposition that biological evolution occurs through the mechanism of natural selection has been almost completely uncontested within the scientific community for much of the 20th century.<ref>An overview of the philosophical, religious, and cosmological controversies by a philosopher who strongly supports evolution is: [[Daniel Dennett]], ''[[Darwin's Dangerous Idea|Darwin's Dangerous Idea: Evolution and the Meanings of Life]]'' (New York: Simon & Schuster, 1995). On the scientific and social reception of evolution in the 19th and early 20th centuries, see: [[Peter J. Bowler]], ''Evolution: The History of an Idea'', 3rd. rev. edn. (Berkeley: University of California Press, 2003).</ref>
Starting with the publication of ''[[The Origin of Species]]'' in 1859, the modern science of evolution has been a source of nearly constant controversy. In general, controversy has centered on the philosophical, [[cosmology|cosmological]], social, and religious implications of evolution, not on the science of evolution itself. The proposition that biological evolution occurs through the mechanism of natural selection has been almost completely uncontested within the scientific community for much of the 20th century.<ref>An overview of the philosophical, religious, and cosmological controversies by a philosopher who strongly supports evolution is: [[Daniel Dennett]], ''[[Darwin's Dangerous Idea|Darwin's Dangerous Idea: Evolution and the Meanings of Life]]'' (New York: Simon & Schuster, 1995). On the scientific and social reception of evolution in the 19th and early 20th centuries, see: [[Peter J. Bowler]], ''Evolution: The History of an Idea'', 3rd. rev. edn. (Berkeley: University of California Press, 2003).</ref>


As Darwin recognized early on, perhaps the most controversial aspect of evolutionary thought is its applicability to human beings. The idea that all diversity in life, including human beings, arose through [[natural science|natural]] processes without a need for supernatural intervention poses difficulties for the [[teleology|belief in purpose]] inherent in most religious faiths &mdash; and especially for the [[Abrahamic religion]]s. Many religious people are able to reconcile the science of evolution with their faith, or see no real conflict<ref>[http://upload.wikimedia.org/wikipedia/commons/1/1d/Evolutionists_Version.sxw]</ref>; [[Judaism]] and [[Evolution and the Roman Catholic Church|Catholicism]] are notable as major faith traditions whose adherents generally see no conflict between evolutionary theory and religious belief.<ref name="RCA">The [[Rabbinical Council of America]] notes that significant Jewish authorities have maintained that evolutionary theory, properly understood, is not incompatible with belief in a Divine Creator, nor with the first 2 chapters of Genesis. [http://www.rabbis.org/news/article.cfm?id=100635]</ref><ref name="Sanhedrin">The [[High Council of B'nei Noah]] a body of non-Jews guided by the Beit Din of B'nei Noah a sub-court of the developing [[Sanhedrin]]: [http://highcouncilofbneinoach.org/resources/Articles/Science_and_Religion.aspx Science and Religion: A proper perspective through an understanding of Hebrew sources] </ref><ref name="aish">[[Aish HaTorah]] [http://www.aish.com/societywork/sciencenature/Age_of_the_universe.asp According to a possible reading of ancient commentators' description of God and nature, the world may be simultaneously young and old.]</ref> The idea that faith and evolution are compatible has been called [[theistic evolution]]. Another group of religious  people, generally referred to as [[creationism|creationists]], consider evolutionary [[origin belief]]s to be incompatible with their faith, their religious texts and [[teleological argument|their perception of design in nature]], and so cannot accept what they call "unguided evolution".
As Darwin recognized early on, perhaps the most controversial aspect of evolutionary thought is its applicability to human beings. The idea that all diversity in life, including human beings, arose through [[natural science|natural]] processes without a need for supernatural intervention poses difficulties for the [[teleology|belief in purpose]] inherent in most religious faiths &mdash; and especially for the [[Abrahamic religion]]s. Many religious people are able to reconcile the science of evolution with their faith, or see no real conflict [[Judaism]] and [[Evolution and the Roman Catholic Church|Catholicism]] are notable as major faith traditions whose adherents generally see no conflict between evolutionary theory and religious belief.<ref name="RCA">The [[Rabbinical Council of America]] notes that significant Jewish authorities have maintained that evolutionary theory, properly understood, is not incompatible with belief in a Divine Creator, nor with the first 2 chapters of Genesis. [http://www.rabbis.org/news/article.cfm?id=100635]</ref><ref name="Sanhedrin">The [[High Council of B'nei Noah]] a body of non-Jews guided by the Beit Din of B'nei Noah a sub-court of the developing [[Sanhedrin]]: [http://highcouncilofbneinoach.org/resources/Articles/Science_and_Religion.aspx Science and Religion: A proper perspective through an understanding of Hebrew sources] </ref><ref name="aish">[[Aish HaTorah]] [http://www.aish.com/societywork/sciencenature/Age_of_the_universe.asp According to a possible reading of ancient commentators' description of God and nature, the world may be simultaneously young and old.]</ref> The idea that faith and evolution are compatible has been called [[theistic evolution]]. Another group of religious  people, generally referred to as [[creationism|creationists]], consider evolutionary [[origin belief]]s to be incompatible with their faith, their religious texts and [[teleological argument|their perception of design in nature]], and so cannot accept what they call "unguided evolution".


One particularly contentious topic evoked by evolution is the biological ''status'' of humanity. Whereas the classical religious view can broadly be characterized as a belief in the [[great chain of being]] (in which people are "above" the animals but slightly "below" the angels), the science of evolution is clear both that humans are animals and that they share common ancestry with [[chimpanzees]], [[gibbon|gibbons]], [[gorillas]], and [[orangutans]]. Some people find the idea of common ancestry repellent, as, in their opinion, it "degrades" humankind. A related conflict arises when critics combine the religious view of people's superior status with the mistaken notion that evolution is necessarily "progressive". If human beings are superior to animals yet evolved from them, these critics claim, "inferior" animals would not still exist. Because animals that are (in their view) "inferior" creatures do demonstrably exist, those criticising evolution sometimes incorrectly take this as supporting their claim that evolution is false.  
One particularly contentious topic evoked by evolution is the biological ''status'' of humanity. Whereas the classical religious view can broadly be characterized as a belief in the [[great chain of being]] (in which people are "above" the animals but slightly "below" the angels), the science of evolution is clear both that humans are animals and that they share common ancestry with [[chimpanzees]], [[gibbon|gibbons]], [[gorillas]], and [[orangutans]]. Some people find the idea of common ancestry repellent, as, in their opinion, it "degrades" humankind. A related conflict arises when critics combine the religious view of people's superior status with the mistaken notion that evolution is necessarily "progressive". If human beings are superior to animals yet evolved from them, these critics claim, "inferior" animals would not still exist. Because animals that are (in their view) "inferior" creatures do demonstrably exist, those criticising evolution sometimes incorrectly take this as supporting their claim that evolution is false.  


In some countries &mdash; notably the [[United States]] &mdash; these and other tensions between religion and science have fueled what has been called the [[creation-evolution controversy]], which, among other things, has generated struggles over the teaching curriculum. While many other fields of science, such as [[physical cosmology|cosmology]] and [[earth science]], also conflict with a literal interpretation of many religious texts, evolutionary studies have borne the brunt of these debates.
In some countries &mdash; notably the USA &mdash; these and other tensions between religion and science have fueled what has been called the [[creation-evolution controversy]], which, among other things, has generated struggles over the teaching curriculum. While many other fields of science, such as [[physical cosmology|cosmology]] and [[earth science]], also conflict with a literal interpretation of many religious texts, evolutionary studies have borne the brunt of these debates.


Evolution has been used to support philosophical and ethical choices which most modern scientists argue are neither mandated by evolution nor supported by science. For example, the [[eugenics|eugenic]] ideas of [[Francis Galton]] were developed into arguments that the human gene pool should be improved by [[selective breeding]] policies, including incentives for reproduction for those of "good stock" and disincentives, such as [[compulsory sterilization]], [[T-4 Euthanasia Program|"euthanasia"]], and later, [[prenatal testing]], [[birth control]], and [[genetic engineering]], for those of "bad". Another example of an extension of evolutionary theory that is widely regarded as unwarranted is "[[Social Darwinism]]"; a term given to the 19th century [[British Whig Party|Whig]] [[Malthusianism|Malthusian]] theory developed by [[Herbert Spencer]] into ideas about "[[survival of the fittest]]" in commerce and human societies as a whole, and by others into claims that [[social inequality]], [[racism]], and [[imperialism]] were justified.<ref>On the history of eugenics and evolution, see [[Daniel Kevles]], ''In the Name of Eugenics: Genetics and the Uses of Human Heredity'' (New York: Knopf, 1985).</ref>
Evolution has been used to support philosophical and ethical choices which most modern scientists argue are neither mandated by evolution nor supported by science. For example, the [[eugenics|eugenic]] ideas of [[Francis Galton]] were developed into arguments that the human gene pool should be improved by [[selective breeding]] policies, including incentives for reproduction for those of "good stock" and disincentives, such as [[compulsory sterilization]], [[T-4 Euthanasia Program|"euthanasia"]], and later, [[prenatal testing]], [[birth control]], and [[genetic engineering]], for those of "bad". Another example of an extension of evolutionary theory that is widely regarded as unwarranted is "[[Social Darwinism]]"; a term given to the 19th century [[British Whig Party|Whig]] [[Malthusianism|Malthusian]] theory developed by [[Herbert Spencer]] into ideas about "[[survival of the fittest]]" in commerce and human societies as a whole, and by others into claims that [[social inequality]], [[racism]], and [[imperialism]] were justified.<ref>On the history of eugenics and evolution, see [[Daniel Kevles]], ''In the Name of Eugenics: Genetics and the Uses of Human Heredity'' (New York: Knopf, 1985).</ref>
Line 306: Line 185:
<references />
<references />
</div>
</div>
===General references===
<div class="references-small" style="column-count:2;-moz-column-count:2;">
*[[Sean B. Carroll]], 2005, ''Endless Forms Most Beautiful: The New Science of Evo Devo and the Making of the Animal Kingdom'', W. W. Norton & Company. ISBN 0-393-06016-0
* {{cite book
| author = Futuyma, D.
|authorlink = Douglas J. Futuyma
| year = 2005
| title = Evolution
| publisher = Sinauer Associates, Inc.
| id = ISBN 0-87893-187-2
}}
*[[Natalia S. Gavrilova]] & [[Leonid A. Gavrilov]], 2002, ''[http://health.families.com/evolution-of-aging-458-467-eoa Evolution of Aging]'', In: David J. Ekerdt (ed.) Encyclopedia of Aging, New York, Macmillan Reference USA, 2002, vol.2, 458-467.ISBN 0-02-865472-2
*Gigerenzer, Gerd, et al., ''The empire of chance: how probability changed science and everyday life'' (New York: Cambridge University Press, 1989).
*Edward J. Larson, ''Evolution: The Remarkable History of a Scientific Theory'' (Modern Library Chronicles). Modern Library (May 4, 2004). ISBN 0-679-64288-9
*Mayr, Ernst. ''What Evolution Is''. Basic Books (October, 2002). ISBN 0-465-04426-3
*Menand, Louis. 2001 ''The Metaphysical Club''. New York: Farar, Straus and Giraux. ISBN 0-374-19963-9
*{{cite journal|author=Smith, D. C.|year=1988|title=Heritable divergence of ''Rhagoletis pomonella'' host races by seasonal asynchrony|journal=Nature|volume=336|pages=66-67|id={{doi|10.1038/336066a0}}|issue=6194}}
*Williams, G.C. (1966). Adaptation and Natural Selection: A Critique of some Current Evolutionary Thought. Princeton, N.J.: Princeton University Press.
*Zimmer, Carl. ''Evolution: The Triumph of an Idea''. Perennial (October 1, 2002). ISBN 0-06-095850-2
*Garcia-Fernàndez, Jordi. [http://www.biolsci.org/v2.php#i3 Amphioxus: a peaceful anchovy fillet to illuminate Chordate Evolution]. Int J Biol Sci (May, 2006).
</div>
==External links==
<!-- IMPORTANT! Please do not add any links before discussing them on the talk page. -->
{{Spoken Wikipedia|Evolution.ogg|2005-04-18}} <!-- updated changed sections 2005-04-18 -->
* [http://www.talkorigins.org Talk.Origins Archive] — see also [[talk.origins]]
* [http://evolution.berkeley.edu/ Understanding Evolution] from [[University of California, Berkeley]]
* [http://nationalacademies.org/evolution/ National Academies Evolution Resources]
* [http://www.evowiki.org/index.php/Main_Page EvoWiki] — A wiki whose goal is to promote general evolution education, and provide mainstream scientific responses to the arguments of antievolutionists.
* [http://www.chains-of-reason.org/chains/evolution-by-natural-selection/introduction.htm Evolution by Natural Selection] — An introduction to the logic of evolution by natural selection
* [http://www.pbs.org/wgbh/evolution/index.html Evolution] — Provided by ''[[Public Broadcasting Service|PBS]]''.
* [http://www.newscientist.com/channel/life/evolution Everything you wanted to know about evolution] — Provided by ''[[New Scientist]]''.
* [http://evol.allenpress.com/evolonline/?request=index-html International Journal of Organic Evolution]
* [http://www.biolsci.org International Journal of Biological Sciences]
* [http://www.necsi.org/projects/evolution/cover/evolution_cover.html New England Complex Systems Institute]
* [http://science.howstuffworks.com/evolution.htm/printable Howstuffworks.com — How Evolution Works]
* [http://anthro.palomar.edu/synthetic/ Synthetic Theory Of Evolution: An Introduction to Modern Evolutionary Concepts and Theories]
* [http://pages.britishlibrary.net/charles.darwin/ Charles Darwin's writings]
* [http://www.genomenewsnetwork.org/categories/index/genome/evolution.php Evolution News from Genome News Network (GNN)]
* [http://www.nap.edu/books/0309063647/html/ National Academy Press: Teaching About Evolution and the Nature of Science]
* [http://www.evolution.mbdojo.com/evolution-for-beginners.html Evolution for beginners]
* [http://www.rmcybernetics.com/science/cybernetics/ai.htm RMCybernetics - AI] Evolution can create emergent behavior in a computer program.
* [http://www.sciencefriday.com/pages/2005/Nov/hour2_111805.html NPR - Science Friday: links to museums, articles and books.]
* [http://www.actionbioscience.org/evolution/lenski.html "Evolution: Fact and Theory" by Richard E. Lenski]
* [http://www.2think.org/evolutionbylevel.shtml Evolution by level] Book reviews of books on evolution by knowledge level.
* [http://www.rationalrevolution.net/articles/understanding_evolution.htm Understanding Evolution: History, Theory, Evidence, and Implications] Deals heavily with the history of evolutionary thought
* [http://www.becominghuman.org/ Becoming Human - Journey through the story of human evolution]
* [http://www.allviewpoints.org/RESOURCES/EVOLUTION/timeline.htm Evolution & ID Timeline] Focuses on major historical and recent events in the scientific and political debate
* [http://www.ucmp.berkeley.edu/history/evotmline.html Timeline of Evolutionary Thought] The life and work of notable people who have contributed to evolutionary thought
;Evolution Simulators
* [http://www.truthtree.com/evolve.shtml Isolated species evolves to interact more efficiently with its environment (java applet)]
* [http://obermuhlner.com/public/Projects/Applets/Blobs/index.html Evolution in a predator-prey relationship (java applet)]
* [http://www.swimbots.com/ Watch small creatures evolve into more efficient swimmers]
* [http://physics.syr.edu/courses/mirror/biomorph/ Blind Watchmaker Applet (java)]
==See also==
:''For a more comprehensive list of topics, see [[:Category:Evolution]] and [[:Category:Evolutionary biology]]''
<p></p>
{| style="background-color: transparent; width: {{{width|100%}}}"
<p></p>
| width="33%" align="{{{align|left}}}" valign="{{{valign|top}}}" |
*[[Abiogenesis]]
*[[Altruism in animals]]
*[[Anagenesis]]
*[[Argument from evolution]]
*[[Atavism]]
*[[Animal evolution]]
*[[Behavioral ecology]]
*[[Catagenesis (biology)|Catagenesis]]
*[[Cladistics]]
*[[Cladogenesis]]
*[[Convergent evolution]]
*[[Creation-evolution controversy]]
*[[Divergent evolution]]
*[[Dual inheritance theory]]
*[[Endosymbiont]]
*[[Eugenics]]
*[[Evolution of sex]]
*[[Evolutionary algorithm]]
*[[Evolutionary art]]
*[[Evolutionary biology]]
<p></p>
| width="33%" align="{{{align|left}}}" valign="{{{valign|top}}}" |
*[[Evolutionary developmental biology]]
*[[Evolutionary medicine]]
*[[Evolution of multicellularity]]
*[[Evolutionary psychology]]
*[[Evolutionary tree]]
*[[Evolutionism]]
*[[Evolvability]]
*[[Experimental evolution]]
*[[Fitness landscape]]
*[[Fossil record]]s
*[[Gene-centered view of evolution]]
*[[Genetic algorithm]]
*[[Genetics]]
*[[Gradualism]]
*[[HeLa]]
*[[Human behavioral ecology]]
*[[Human evolution]]
*[[Instinct]]
*[[Kin selection]]
*[[Language]]
<p></p>
| width="33%" align="{{{align|left}}}" valign="{{{valign|top}}}" |
*[[List of publications on evolution and human behavior]]
*[[Modern evolutionary synthesis]]
*[[Natural science]]
*[[Natural selection]]
*[[Neutral theory of molecular evolution]]
*[[Niche construction]]
*[[Origin of life]]
*[[Paleontology]]
*[[Parallel evolution]]
*[[Parental investment]]
*[[Punctuated equilibrium]]
*[[Quantum evolution]]
*[[Quasispecies model]]
*[[Reciprocal altruism]]
*[[Scientific method]]
*[[Sexual selection]]
*[[Social effect of evolutionary theory]]
*[[Sociobiology]]
*[[Teratogenesis]]
*[[Timeline of evolution]]
<p></p>
|}
{{evolution}}
{{popgen}}
{{featured article}}
<!-- Categorization -->
<!-- Localization -->
[[Category:Evolutionary biology]]
[[Category:Evolution| ]]
[[Category:Theories]]
[[af:Evolusie]]
[[ar:نظرية النشوء]]
[[zh-min-nan:Chìn-hoà]]
[[bn:বিবর্তন]]
[[bg:Еволюция]]
[[ca:Teoria de l'evolució]]
[[cs:Evoluce]]
[[cy:Esblygiad]]
[[da:Evolution]]
[[de:Biologische Evolution]]
[[es:Evolución biológica]]
[[eo:Evoluismo]]
[[fa:فرگشت]]
[[fr:Évolution]]
[[ko:진화]]
[[id:Evolusi]]
[[it:Evoluzione]]
[[he:אבולוציה]]
[[lb:Evolutioun]]
[[lt:Evoliucija]]
[[hu:Evolúció]]
[[mk:Еволуција]]
[[nl:Evolutietheorie]]
[[ja:進化]]
[[no:Evolusjon]]
[[pl:Ewolucja biologiczna]]
[[pt:Evolução]]
[[ro:Teoria evoluţionistă]]
[[ru:Эволюционное учение]]
[[sk:Evolúcia]]
[[sl:Evolucija]]
[[su:Évolusi]]
[[fi:Evoluutio]]
[[sv:Evolution]]
[[th:วิวัฒนาการ]]
[[tr:Evrim]]
[[yi:עוועלאציע]]
[[zh:进化论]]
{{CZ live}}

Latest revision as of 16:55, 24 May 2012

This article is developing and not approved.
Main Article
Discussion
Related Articles  [?]
Bibliography  [?]
External Links  [?]
Citable Version  [?]
Video [?]
 
This editable Main Article is under development and subject to a disclaimer.

In biology, the concept of evolution applies to a natural process whereby a distinct population of living systems, a biological species, for example, changes in its biological characteristics over generations, often with many new and diverse types of living systems emerging in the process. That application of the word 'evolution' to living systems does not exhaust the biological systems the word applies to; it includes transgenerational molecular evolution and language evolution, for example.

In his 1998 edition of Evolutionary Biology, evolutionary biologist Douglas Futuyma defined 'evolution' thus:

"In its broadest sense, evolution simply means "change"...Usually..we don't apply the term to the changes an individual entity undergoes during its lifetime [i.e., its 'ontogeny']. Rather, an evolving system is ordinarily one in which there is descent of entities [viz., populations of similar entities], one generation from another, over time, and in which characteristics of the entities differ across generations. Thus evolution in a broad sense is descent with modification, and often with diversification."[1]

Expanding in his 2009 edition of Evolution, he writes:

"It [‘evolution’] is sometimes used to describe changes in individual objects such as stars. Biological' (or organic) evolution, however, is change in the properties of groups of organisms over the course of generations. The development, or ONTOGENY, of an individual organism is not consid¬ered evolution: individual organisms do not evolve. Groups of organisms, which we may call populations, undergo descent with modification. Populations may become subdivided, so that several populations are derived from a common ancestral population. If different changes transpire in the several populations, the populations diverge.”[2]

In one of the specific forms the process of evolution unrolls, a closely related group of organisms, referred to as a 'population', undergoes modifications of its characteristics—traits, or so-called phenotypes—over a period of generations in a manner that subdivides a parent population into distinctly differing descendant populations, a process in which a common ancestral population gives rise to diverse populations—the process of change referred to as 'descent with modification'.

During the 1800s, before mid-century, two factors in combination had made biological evolution plausible and likely to open-minded leading scientists, such as the eminent geologist, Charles Lyell[3], and biologist Alfred Russel Wallace. The first: Evidence from strategraphic geology suggested a much older age of the earth than the approximately 4,000 years biblical scholars had determined for the Christian orthodox view, important evidence because of a constraining power of Christianity on the acceptance of scientific ideas that contradicted accepted interpretation of the Christian bible. No less important, the much older age of the earth would have extended greatly a time for biological evolution to occur. The second: Evidence from paleontology indicated the existence of unfamiliar species from the time of the old earth, in the old geographic strata, now extinct.

Thus, biological evolution was plausible and considered likely in science by the mid-1800s because the earth appeared very old and because species existed in the old earth, but not the contemporary species, extinct forms from which contemporary species might have descended by transformation. By mid-century the time was ripe for theories to explain how a descendant evolution might occur, and for a few, ripe for strictly naturalistic theories.

The word 'evolution' derives from the Latin word evolvere, which translates to 'to unfold', or 'to unroll', as if to unroll an ancient scroll and display the intellectual potentialities hidden within, or to unfold the map of time-development of biological diversity.[4] Before the word 'evolution' was introduced in biology to refer to the process of transgenerational changes in species, the process was referred to as 'transmutation', or 'transformation', the concepts, 'transmutationism' and 'transformationism'.[5]

As mentioned above, 'evolution' applies not only to biological evolution, the topic of the present article, but also to other domains of nature that change in a descendant way similar to that of biological evolution, such as the evolution of the universe itself.[6]


Among the many ways biologists recognize the explanatory power of the concept of evolution, they can account for the vast biodiversity of living systems on Earth, including the many extinct species attested in the fossil record. An evolutionary perspective can lead to insights for preventing, diagnosing and treating human diseases.

Most biologists endorse the proposition of the 20th century's pioneering evolutionary biologist, Theodosius Dobzhansky, videlicet: "Nothing in biology makes sense except in the light of evolution."[7]  For evolutionary biologists, not surprisingly, evolution takes center stage, in particular as "the unifying theory of biology".[1] [8]

Biologists distinguish between the factuality of evolution and theories of its mechanism.

Although there is overwhelming scientific consensus supporting the reality of biological evolution, because of its potential implications for the origins of humankind, evolutionary theory has been at the center of many social and religious controversies since its inception.

Brief history of the idea of evolution

The concept that plants and animals change form over generations has existed since ancient times, notably among Greek philosophers such as Anaximander[9] and Epicurus[10] [11] and Indian philosophers such as Patañjali.

Evolution is not so much a modern discovery as some of its advocates would have us believe. It made its appearance early in Greek philosophy, and maintained its position more or less, with the most diverse modifications, and frequently confused with the idea of emanation, until the close of ancient thought. The Greeks had, it is true, no term exactly equivalent to ” evolution”; but when Thales asserts that all things originated from water; when Anaximenes calls air the principle of all things, regarding the subsequent process as a thinning or thickening, they must have considered individual beings and the phenomenal world as, a result of evolution, even if they did not carry the process out in detail. Anaximander is often regarded as a precursor of the modem theory of development. He deduces living beings, in a gradual development, from moisture under the influence of warmth, and suggests the view that men originated from animals of another sort, since if they had come into existence as human beings, needing fostering care for a long time, they would not have been able to maintain their existence. In Empedocles, as in Epicurus and Lucretius, who follow in Hs [sic] footsteps, there are rudimentary suggestions of the Darwinian theory in its broader sense; and here too, as with Darwin, the mechanical principle comes in; the process is adapted to a certain end by a sort of natural selection, without regarding nature as deliberately forming its results for these ends. [9]

The ancient Greeks started the naturalists and devotees of natural history, as well as the world of science known as natural philosophy, on the path of thinking about change in the animate and inanimate world and the implications of those thoughts on the changes in the recent past and on future change. By the mid-1400s ideas of evolution began fast growing in abundance. Them concept of evolution responded to the fuel of expanding knowledge of fossilized living things, and the recognition of Earth's history, its long time of existence, coming up with the idea that the available time favored evolution of present living things from ancestors that looked different in their fossilized remains.

Mechanisms that mediate evolution

Jean-Baptiste Lamarck theorized that living things passed on their traits to offspring, but his theories differed from modern evolutionary theory in that he believed that acquired traits were passed on. Thus, Lamarckian evolution posits that the giraffe acquired its long neck from stretching to reach higher foliage. An individual giraffes lengthened its neck from a lifetime of stretching, and passed on the longer neck to its offspring, which themselves stretched to reach even-higher branches, creating a cycle that concluded with the long necks of today's giraffes.[12]

The transmutation of species was accepted by many scientists before 1859, but Charles Darwin's On The Origin of Species by Means of Natural Selection provided the first convincing exposition[13] of a mechanism by which evolutionary change could occur: natural selection. After many years of working in private on his theory, Darwin was motivated to publish his work on evolution when he received a letter from Alfred Russell Wallace in which Wallace revealed his own, independent discovery of natural selection. Accordingly, Wallace is usually given shared credit for originating the theory.[14]

Darwin's book sparked a great deal of scientific and social debate. Darwin's work relied on many different fields of scientific inquiry for its evidence, and debates over the theory took place in many different arenas. Although the occurrence of biological evolution was generally accepted by scientists, Darwin's specific ideas about evolution—that it occurred gradually, through natural selection—were contested. Additionally, while Darwin was able to observe variation, and infer natural selection and thereby adaptation, he was unable to explain how variation might arise, or be altered over generations. Darwin's proposal of a hereditary mechanism (pangenesis) lacked evidence and was ultimately rejected,[15] being replaced by genetics.

From the end of the 19th century through the early 20th century, forms of neo-Lamarckism, "progressive" evolution (orthogenesis), and an evolution which worked by "jumps" (saltationism, as opposed to gradualism) became popular, although a form of neo-Darwinism, led by August Weismann, also enjoyed some minor success. The biometric school of evolutionary theory, resulting from the work of Darwin's cousin, Francis Galton, emerged as well, using statistical approaches to biology which emphasized gradualism and some aspects of natural selection.[16]

Mendel's work was "rediscovered" in 1901. Debates over various aspects of how evolution occurs have continued. One prominent debate was over the theory of punctuated equilibrium, proposed in 1972 by paleontologists Niles Eldredge and Stephen Jay Gould to explain the paucity of gradual transitions between species in the fossil record, as well as the absence of change or stasis that is observed over significant intervals of time.

Academic disciplines

Evolutionary biology is concerned with the origin and descent of species, as well as their changes over time. It is an interdisciplinary field including scientists from many traditional taxonomically-oriented disciplines, including scientists with specialist training in particular organisms, such as mammalogy, ornithology, or herpetology, but who use those organisms to answer general questions in evolution. Evolutionary biology as an academic discipline in its own right emerged as a result of the modern evolutionary synthesis in the 1930s and 1940s.

Evolutionary developmental biology (informally, evo-devo) is a field of biology that compares the developmental processes of different animals in an attempt to determine the ancestral relationship between organisms and how developmental processes evolved. The discovery of genes regulating development in model organisms allowed for comparisons to be made with genes and genetic networks of related organisms.

Physical anthropology emerged in the late 19th century as the study of human osteology, and the fossilized skeletal remains of other hominids. At that time, anthropologists debated whether their evidence supported Darwin's claims, because skeletal remains revealed temporal and spatial variation among hominids, but Darwin had not offered an explanation of the specific mechanisms that produce variation. With the recognition of Mendelian genetics and the rise of the modern synthesis, however, evolution became both the fundamental conceptual framework for, and the object of study of, physical anthropologists. In addition to studying skeletal remains, they began to study genetic variation among human populations (population genetics); thus, some physical anthropologists began calling themselves biological anthropologists.

Fossil Record

Fossils are critical evidence for estimating when various lineages originated. Since fossilization of an organism is an uncommon occurrence, usually requiring hard parts (like teeth, bone or pollen), the fossil record is traditionally thought to provide only sparse and intermittent information about ancestral lineages. Fossilization of organisms without hard body parts is rare, but does happen under unusual circumstance; very rapid burial, and both low oxygen environment and sparse microbial action[17].

The fossil record contains the earliest known examples of life itself, as well as the earliest occurrences of individual lineages. The first complex animals date from the early Cambrian period, approximately 520 million years ago. The records of individual species yield information regarding the patterns and rates of evolution, showing for example if species evolve into new species (speciation) gradually and incrementally, or in relatively brief intervals of geologic time. The fossil record is a document of large scale patterns and events in the history of life, many of which have influenced the evolutionary history of numerous lineages. Mass extinctions frequently resulted in the loss of entire groups of species, such as the non-avian dinosaurs, while leaving others relatively unscathed. Recently, molecular biologists have used the time since divergence of related lineages to calibrate the rate at which mutations accumulate, and at which the genomes of different lineages evolve.

Phylogenetics, the study of the ancestry of species, has revealed that structures with similar internal organization may perform divergent functions. Vertebrate limbs are a common example of such homologous structures. The appendages on bat wings, for example, are very structurally similar to human hands, and may constitute a vestigial structure. Other examples include the presence of hip bones in whales and snakes. Such structures may exist with little or no function in a more current organism, yet have a clear function in an ancestral species of the same. Examples of vestigial structures in humans include wisdom teeth, the coccyx and the vermiform appendix.

Molecular evidence

Comparison of DNA sequences allows organisms to be grouped by sequence similarity, and the resulting phylogenetic trees are typically congruent with traditional taxonomy, and are often used to strengthen or correct taxonomic classifications. Sequence comparison is considered a measure robust enough to be used to correct erroneous assumptions in the phylogenetic tree in instances where other evidence is scarce. For example, neutral human DNA sequences are approximately 1.2% divergent (based on substitutions) from those of their nearest genetic relative, the chimpanzee, 1.6% from gorillas, and 6.6% from baboons.[18] Genetic sequence evidence thus allows inference and quantification of genetic relatedness between humans and other apes.[19][20] The sequence of the 16S rRNA gene, a vital gene encoding a part of the ribosome, was used to find the broad phylogenetic relationships between all extant life. The analysis, originally done by Carl Woese, resulted in the three-domain system, arguing for two major splits in the early evolution of life. The first split led to modern Bacteria and the subsequent split led to modern Archaea and Eukaryote.

The proteomic evidence also supports the universal ancestry of life. Vital proteins, such as the ribosome, DNA polymerase, and RNA polymerase are found in the most primitive bacteria to the most complex mammals. The core part of the protein is conserved across all lineages of life, serving similar functions. Higher organisms have evolved additional protein subunits, largely affecting the regulation and protein-protein interaction of the core. Other overarching similarities between all lineages of extant organisms, such as DNA, RNA, amino acids, and the lipid bilayer, give support to the theory of common descent. The chirality of DNA, RNA, and amino acids is conserved across all known life. As there is no functional advantage to right or left handed molecular chirality, the simplest hypothesis is that the choice was made randomly in the early beginnings of life and passed on to all extant life through common descent.

Molecular evidence also offers a mechanism for large evolutionary leaps and macroevolution. Horizontal gene transfer, the process in which an organism transfers genetic material (i.e. DNA) to another cell that is not its offspring, allows for large sudden evolutionary leaps in a species by incorporating beneficial genes evolved in another species. The Endosymbiotic theory explains the origin of mitochondria and plastids (e.g. chloroplasts), which are organelles of eukaryotic cells, as the incorporation of an ancient prokaryotic cell into ancient eukaryotic cell. Rather than evolving eukaryotic organelles slowly, this theory offers a mechanism for a sudden evolutionary leap by incorporating the genetic material and biochemical composition of a separate species. This evolutionary mechanism has been observed. Heneta, a protist, is an extant organism that is undergoing endosymbiotic evolution.[21][22]

Further evidence for reconstructing ancestral lineages comes from so-called junk DNA such as pseudogenes, i.e., 'dead' genes, which steadily accumulate mutations.[23]

Mutation

What accounts for genetic variation? Mutations are permanent, transmissible changes to the genetic material of a cell. These changes can be caused by: "copying errors" in the genetic material during cell division. (Explain frame shift) Exposure to radiation, certain chemicals, and some viruses can destabilize these genetic molecules and cause biochemical changes that are permanent. When mutations are made in the genes of a cell they ordinarily will be passed on to daughter cells when the cell divides. In multicellular organisms, the mutations will be passed on to progeny if germline mutations, that is- if the genes in the gametes are changed. Mutations that are not affected by natural selection are called neutral mutations. Their frequency in the population is governed by mutation rate, genetic drift and selective pressure on linked alleles. It is understood that most of a species' genome, in the absence of selection, undergoes a steady accumulation of neutral mutations.

Mobile elements, transposons, make up a major fraction of the genomes of plants and animals and appear to have played a significant role in the evolution of genomes. These mobile insertional elements can jump within a genome and alter existing genes and gene networks to produce evolutionary change and diversity.

Selection and adaptation

For more information, see: Natural selection and Adaptation.


Natural selection can be subdivided into two categories:

  • Ecological selection occurs when organisms that survive and reproduce increase the frequency of their genes in the gene pool over those that do not survive.
  • Sexual selection occurs when organisms which are more attractive to the opposite sex because of their features reproduce more and thus increase the frequency of those features in the gene pool.

Natural selection also operates on mutations in several different ways:

  • Positive or directional selection increases the frequency of a beneficial mutation, or pushes the mean in either direction.
  • Purifying or stabilizing selection maintains a common trait in the population by decreasing the frequency of harmful mutations and weeding them out of the population. "Living fossils" are arguably the product of stabilizing selection, as their form and traits have remained virtually identical over a long period of time. It is argued that stabilizing selection is the most common form of natural selection.
  • Artificial selection refers to purposeful breeding of a species to produce a more desirable and “perfect” breed. Humans have directed artificial selection in the breeding of both animals and plants, with examples ranging from agriculture (crops and livestock) to pets and horticulture. However, because humans are only part of the environment, the fractions of change in a species due to natural or artificial means can be difficult to determine. Artificial selection within human populations is a controversial enterprise known as eugenics.
  • Balancing selection maintains variation within a population through a number of mechanisms, including:
  • Disruptive selection favors both extremes, and results in a bimodal distribution of gene frequency. The mean may or may not shift.
  • Selective sweeps describe the affect of selection acting on linked alleles. It comes in two forms:
    • Background selection occurs when a deleterious mutation is selected against, and linked mutations are eliminated along with the deleterious variant, resulting in lower genetic polymorphism in the surrounding region.
    • Genetic hitchhiking occurs when a beneficial allele is selected for, and linked alleles, which can be neutral or beneficial, are pushed towards fixation along with the beneficial allele.


Most biologists believe that adaptation occurs through the accumulation of many mutations of small effect. However, macromutation is an alternative process for adaptation that involves a single, very large scale mutation.

Recombination

In asexual organisms, variants in genes on the same chromosome will always be inherited together - they are linked, by virtue of being on the same DNA molecule. However, sexual organisms, in the production of gametes, shuffle linked alleles on homologous chromosomes inherited from the parents via meiotic recombination. This shuffling allows independent assortment of alleles (mutations) in genes to be propagated in the population independently. This allows bad mutations to be purged and beneficial mutations to be retained more efficiently than in asexual populations.

However, the meitoic recombination rate is not very high - on the order of one crossover (recombination event between homomolgous chromosomes) per chromosome arm per generation. Therefore, linked alleles are not perfectly shuffled away from each other, but tend to be inherited together. This tendency may be measured by comparing the co-occurrence of two alleles, usually quantified as linkage disequilibrium (LD). A set of alleles that are often co-propagated is called a haplotype. Strong haplotype blocks can be a product of strong positive selection.

Gene flow and population structure

Gene flow (also called gene admixture or simply migration) is the exchange of genetic variation between populations, when geography and culture are not obstacles. Ernst Mayr thought that gene flow is likely to be homogenising, and therefore counteract selective adaptation. Where there are obstacles to gene flow, the situation is termed reproductive isolation and is considered to be necessary for speciation.

The free movement of alleles through a population may also be impeded by population structure. For example, most real-world populations are not actually fully interbreeding; geographic proximity has a strong influence on the movement of alleles within the population.

An example of the effect of population structure is the so-called founder effect, resulting from a migration or population bottleneck, in which a population temporarily has very few individuals, and therefore loses a lot of genetic variation. In this case, a single, rare allele may suddenly increase very rapidly in frequency within a specific population if it happened to be prevalent in a small number of "founder" individuals. The frequency of the allele in the resulting population can be much higher than otherwise expected, especially for deleterious, disease-causing alleles. Since population size has a profound effect on the relative strengths of genetic drift and natural selection, changes in population size can alter the dynamics of these processes considerably.

Drift

Genetic drift describes changes in allele frequency from one generation to the next due to sampling variance. The frequency of an allele in the offspring generation will vary according to a probability distribution of the frequency of the allele in the parent generation. Thus, over time even in the absence of selection upon the alleles, allele frequencies will tend to "drift" upward or downward, eventually becoming "fixed" - that is, going to 0% or 100% frequency. Thus, fluctuations in allele frequency between successive generations may result in some alleles disappearing from the population due to chance alone. Two separate populations that begin with the same allele frequencies therefore might drift apart by random fluctuation into two divergent populations with different allele sets (for example, alleles present in one population could be absent in the other, or vice versa).

The impact of genetic drift depends strongly on the size of the population (generally abbreviated as N): drift is important in small mating populations (see Founder effect and Population bottleneck), where chance fluctuations from generation to generation can be large. The relative importance of natural selection and genetic drift in determining the fate of new mutations also depends on the population size and the strength of selection: when N times s (population size times strength of selection) is small, genetic drift predominates. When N times s is large, selection predominates. Thus, natural selection is predominant in large populations, or equivalently, genetic drift is stronger in small populations. Finally, the time for an allele to become fixed in the population by genetic drift (that is, for all individuals in the population to carry that allele) depends on population size, with smaller populations requiring a shorter time to fixation.

Speciation and extinction

Speciation is the process by which new biological species arise. This may take place by various mechanisms. Allopatric speciation occurs in populations that become isolated geographically, such as by habitat fragmentation or migration.[24]. Sympatric speciation occurs when new species emerge in the same geographic area[25][26] Ernst Mayr's peripatric speciation is a type of speciation that exists in between the extremes of allopatry and sympatry. Peripatric speciation is a critical underpinning of the theory of punctuated equilibrium. An example of rapid sympatric speciation can be eloquently represented in the triangle of U; where new species of Brassica sp. have been made by the fusing of separate genomes from related plants.

Extinction is the disappearance of species (i.e. gene pools). The moment of extinction generally occurs at the death of the last individual of that species. Extinction is not an unusual event in geological time — species are created by speciation, and disappear through extinction. The Permian-Triassic extinction event was the Earth's most severe extinction event, rendering extinct 90% of all marine species and 70% of terrestrial vertebrate species. In the Cretaceous-Tertiary extinction event many forms of life perished (including approximately 50% of all genera), the most often mentioned among them being the extinction of the non-avian dinosaurs.

Current research

Evolution is still an active field of research in the scientific community. Improvements in sequencing methods have resulted in a large increase of sequenced genomes, allowing for the testing and refining of the theory of evolution with respect to whole genome data. Advances in computational hardware and software have allowed for the testing and extrapolation of increasingly advanced evolutionary models. Discoveries in biotechnology have produced methods for the ‘’de novo’’ synthesis of proteins and, potentially, entire genomes, driving evolutionary studies at the molecular level.

Common misunderstandings about evolutionary theory

Progressiveness, complexity, and evolution

Evolutionary processes and, in general, scientific explanations of the world are often in contrast with the immediate and simple explanations that our brain gives of reality (e.g. the sun seems to turn around the earth, the earth seems to be flat), and are influenced by what Francis Bacon called "idola"[[27]] (false notions or tendencies which distort the truth [[28]]).[29]

One of the most common misunderstandings of evolution is that one species can be "more highly evolved" than another, that evolution is necessarily progressive and/or leads to greater "complexity", or that its converse is "devolution".[30] Evolution provides no assurance that later generations are more intelligent or complex than earlier generations. The claim that evolution results in progress is not part of modern evolutionary theory; it derives from earlier belief systems which were held around the time Darwin devised his theory of evolution.

Evolution has involved "progression" towards more complexity, since all of the earliest lifeforms were extremely simple and there was nowhere to go but up. However, there is no guarantee that any particular organism existing today will become more intelligent, more complex, bigger, or stronger in the future. In fact, natural selection will only favor this kind of "progression" if it has a variant to act upon that increases chance of survival, i.e. the ability to live long enough to raise offspring to sexual maturity. The same mechanism can actually favor lower intelligence, lower complexity, and so on if those traits become a selective advantage in the organism's environment.

One way of understanding the apparent "progression" of lifeforms over time is to remember that the earliest life began as components of cells. Evolution caused life to become more complex, since becoming simpler wasn't advantageous. Once individual lineages have attained sufficient complexity, simplifications (specialization) become more likely.

Speciation

It is sometimes claimed that speciation – the origin of new species – has never been directly observed, and thus evolution cannot be called sound science. This is a misunderstanding of both science and evolution. First, scientific discovery does not occur solely through reproducible experiments; the principle of uniformitarianism allows natural scientists to infer causes through their empirical effects. Moreover, since the publication of On the Origin of Species scientists have confirmed Darwin's hypothesis by data gathered from sources that did not exist in his day, such as DNA similarity among species and new fossil discoveries. Finally, speciation has actually been directly observed.[31] (See the hawthorn fly example.)

A variation of this assertion that microevolution has been observed and macroevolution has not been observed is subject to the same counterarguments. While there is debate over the details of how evolution happened, it is generally accepted that macroevolution uses the same mechanisms of change as those already observed in microevolution.

Purposiveness and evoluton

Peter Bowler, eminent historian of science, in his 3rd edition of his widely cited Evolution: History of an Idea, writes:

The antagonism of the creationists should not blind us to the fact that science and religion have sometimes been able to work in harmony. The history of evolutionism reveals many attempts to see the development of life on earth as the unfolding of a divine plan. There have certainly been some scientists who would adopt a more militant posture, arguing that humanity simply has to come to terms with the unpleasant fact that it is the product of a purposeless sequence of natural events.[14]

Yes, we humans may have resulted from “a purposeless sequence of natural events”, but the fact of that happening to us I do not find “unpleasant”. The purposelessness of the sequence of natural events in evolution, even considering biological evolution only, does not necessarily exclude, in a lineage of entities, that new kinds of evolvable entities might emerge that could impose purpose on evolutionary change. Indeed, human culture emerged, in part from new entities called ‘memes’ or ‘culturgens’, whose evolvable complex associations create knowledge and ideas. The eugenics movement early exemplified the potentiality for the lineage culminating in Homo sapiens to purposefully order the sequence of natural events.

Social and religious controversies

Starting with the publication of The Origin of Species in 1859, the modern science of evolution has been a source of nearly constant controversy. In general, controversy has centered on the philosophical, cosmological, social, and religious implications of evolution, not on the science of evolution itself. The proposition that biological evolution occurs through the mechanism of natural selection has been almost completely uncontested within the scientific community for much of the 20th century.[32]

As Darwin recognized early on, perhaps the most controversial aspect of evolutionary thought is its applicability to human beings. The idea that all diversity in life, including human beings, arose through natural processes without a need for supernatural intervention poses difficulties for the belief in purpose inherent in most religious faiths — and especially for the Abrahamic religions. Many religious people are able to reconcile the science of evolution with their faith, or see no real conflict Judaism and Catholicism are notable as major faith traditions whose adherents generally see no conflict between evolutionary theory and religious belief.[33][34][35] The idea that faith and evolution are compatible has been called theistic evolution. Another group of religious people, generally referred to as creationists, consider evolutionary origin beliefs to be incompatible with their faith, their religious texts and their perception of design in nature, and so cannot accept what they call "unguided evolution".

One particularly contentious topic evoked by evolution is the biological status of humanity. Whereas the classical religious view can broadly be characterized as a belief in the great chain of being (in which people are "above" the animals but slightly "below" the angels), the science of evolution is clear both that humans are animals and that they share common ancestry with chimpanzees, gibbons, gorillas, and orangutans. Some people find the idea of common ancestry repellent, as, in their opinion, it "degrades" humankind. A related conflict arises when critics combine the religious view of people's superior status with the mistaken notion that evolution is necessarily "progressive". If human beings are superior to animals yet evolved from them, these critics claim, "inferior" animals would not still exist. Because animals that are (in their view) "inferior" creatures do demonstrably exist, those criticising evolution sometimes incorrectly take this as supporting their claim that evolution is false.

In some countries — notably the USA — these and other tensions between religion and science have fueled what has been called the creation-evolution controversy, which, among other things, has generated struggles over the teaching curriculum. While many other fields of science, such as cosmology and earth science, also conflict with a literal interpretation of many religious texts, evolutionary studies have borne the brunt of these debates.

Evolution has been used to support philosophical and ethical choices which most modern scientists argue are neither mandated by evolution nor supported by science. For example, the eugenic ideas of Francis Galton were developed into arguments that the human gene pool should be improved by selective breeding policies, including incentives for reproduction for those of "good stock" and disincentives, such as compulsory sterilization, "euthanasia", and later, prenatal testing, birth control, and genetic engineering, for those of "bad". Another example of an extension of evolutionary theory that is widely regarded as unwarranted is "Social Darwinism"; a term given to the 19th century Whig Malthusian theory developed by Herbert Spencer into ideas about "survival of the fittest" in commerce and human societies as a whole, and by others into claims that social inequality, racism, and imperialism were justified.[36]

References

  1. 1.0 1.1 Futuyma DJ. (1998) Evolutionary Biology 3rd ed. Sinauer Associates, Inc., Sunderland MA ISBN 0-87893-189-9
  2. Futuyma DJ. (2009) Evolution. 2nd ed. Sunderland, MA: Sinauer Associates. ISBN 978-0-87893-223-8.
  3. Principles of Geology
  4. Futuyma DJ. (2005) Evolution. 1st ed. Sunderland, Mass.: Sinauer Associates. ISBN 0878931872. | Downloadable Chapter 15 2nd. ed. "Sex and Reproductive Success".
  5. Wilkins J. (1997) Darwin's Precursors and Influences: Introduction
  6. Smolin L. (2008) The Life of the Cosmos. Oxford University Press. ISBN 9780199216802. | Google Books preview.
  7. Dobzhansky T (1973). "Nothing in biology makes sense except in the light of evolution". Am Biol Teach: 125-9.
  8. Introduction to Evolutionary Biology
  9. 9.0 9.1 History of Evolution. Internet Encyclopedia of Philosophy.
  10. Konstan D. (2009) "Epicurus", [http://plato.stanford.edu/archives/spr2009/entries/epicurus/ The Stanford Encyclopedia of Philosophy (Spring 2009 Edition), Edward N. Zalta (ed.)
    • "The philosophy of Epicurus (341–270 B.C.) was a complete and interdependent system, involving a view of the goal of human life (happiness, resulting from absence of physical pain and mental disturbance), an empiricist theory of knowledge (sensations, including the perception of pleasure and pain, are infallible criteria), a description of nature based on atomistic materialism, and a naturalistic account of evolution, from the formation of the world to the emergence of human societies." [emphasis added]
  11. Evolution - Epicurus Wiki
  12. Early Concepts of Evolution: Jean Baptiste Lamarck
  13. In the years after Darwin's publication, numerous "predecessors" to natural selection were discovered, such as William Charles Wells and Patrick Matthew, who had published unelaborated and undeveloped versions of similar theories earlier to little or no attention. Darwin was the first to develop the theory rigorously and developed it independently. On Matthew, one historian of evolution has written that he "did suggest a basic idea of selection, but he did nothing to develop it; and he published it in the appendix to a book on the raising of trees for shipbuilding. No one took him seriously, and he played no role in the emergence of Darwinism. Simple priority is not enough to earn a thinker a place in the history of science: one has to develop the idea and convince others of its value to make a real contribution. Darwin's notebooks confirm that he drew no inspiration from Matthew or any of the other alleged precursors." Bowler, PJ (2003). Evolution: The History of an Idea. Berkeley: University of California Press, 158. 
  14. 14.0 14.1 Bowler,P.J. (2003) Evolution: The History of an Idea. 3rd ed., completely rev. and expanded. Berkeley: University of California Press. ISBN 052236939. | Google Books preview. $From chapter 1: "Much can be gained from the lessons of history, especially an awareness of the dangers of oversimplification. Before we can undertake such a study, however, we must establish a framework for understanding the rise of modern evolutionism. We must identify the full range of issues over which evolutionism has challenged the traditional worldview. We must appreciate how aspects of the old view can be modernized to fit in with certain kinds of evolutionary thought. The basic idea of evolution can be developed in many different ways, each with its own broader implications. We must also look more closely at the problems facing the historian who has to seek a balance between the conventional image of science as the objective evaluation of factual data and the suspicion of many critics that scientific theories are value-laden contributions to philosophical and ideological debates."
  15. Darwin’s Theory of Pangenesis
  16. Bowler, Peter J. (1989). The Mendelian Revolution: The Emergence of Hereditarian Concepts in Modern Science and Society. Baltimore: John Hopkins University Press. 
  17. Schweitzer M.H. et al (2005). "Soft-tissue vessels and cellular preservation in Tyrannosaurus rex". Science 307 (5717): 1952-1955.
  18. Two sources: 'Genomic divergences between humans and other hominoids and the effective population size of the common ancestor of humans and chimpanzees'. and 'Quantitative Estimates of Sequence Divergence for Comparative Analyses of Mammalian Genomes' "[1] [2]"
  19. The picture labeled "Human Chromosome 2 and its analogs in the apes" in the article Comparison of the Human and Great Ape Chromosomes as Evidence for Common Ancestry is literally a picture of a link in humans that links two separate chromosomes in the nonhuman apes creating a single chromosome in humans. It is considered a missing link, and the ape-human connection is of particular interest. Also, while the term originally referred to fossil evidence, this too is a trace from the past corresponding to some living beings which when alive were the physical embodiment of this link.
  20. The New York Times report Still Evolving, Human Genes Tell New Story, based on A Map of Recent Positive Selection in the Human Genome, states the International HapMap Project is "providing the strongest evidence yet that humans are still evolving" and details some of that evidence.
  21. Okamoto N, Inouye I. (2005). "A secondary symbiosis in progress". Science 310 (5746): 287.
  22. Okamoto N, Inouye I (2006). "Hatena arenicola gen. et sp. nov., a Katablepharid Undergoing Probable Plastid Acquisition.". Protist.
  23. Pseudogene evolution and natural selection for a compact genome. "[3]"
  24. Hoskin et al (Oct 2005). "Reinforcement drives rapid allopatric speciation". Nature 437: 1353-1356.
  25. Savolainen et al (May 2006). "Sympatric speciation in palms on an oceanic island". Nature 441: 210-213.
  26. Barluenga et al (February 2006). "Sympatric speciation in Nicaraguan crater lake cichlid fish". Nature 439: 719-723.
  27. Hall MP. The Four Idols of Francis Bacon: The New Instrument of Knowledge.
    • "In the Novum Organum (the new instrumentality for the acquisition of knowledge) Francis Bacon classified the intellectual fallacies of his time under four headings which he called idols. He distinguished them as idols of the Tribe, idols of the Cave, idols of the Marketplace and idols of the Theater…An idol is an image, in this case held in the mind, which receives veneration but is without substance in itself. Bacon did not regard idols as symbols, but rather as fixations."
  28. Fantini F. (2005) Didattica dell'evoluzione. In Evoluzione tra ricerca e didattica, XIV – Special number Edited by: Associazione Nazionale Insegnanti di Scienze Naturali. Agnano Pisano: Stamperia Editoriale Pisana; 2005:203-209.
  29. Guidetti R, Baraldi L, Calzolai C, Pini L, Veronesi P, Pederzoli A. (2007) Fantastic animals as an experimental model to teach animal adaptation. BMC Evolutionary Biology 7(Suppl 2):S13 doi: 10.1186/1471-2148-7-S2-S13.
  30. talkorigins Claim CB932: Evolution of degenerate forms
  31. Boxhorn, Joseph. Observed Instances of Speciation. Talk Origins Archive.
  32. An overview of the philosophical, religious, and cosmological controversies by a philosopher who strongly supports evolution is: Daniel Dennett, Darwin's Dangerous Idea: Evolution and the Meanings of Life (New York: Simon & Schuster, 1995). On the scientific and social reception of evolution in the 19th and early 20th centuries, see: Peter J. Bowler, Evolution: The History of an Idea, 3rd. rev. edn. (Berkeley: University of California Press, 2003).
  33. The Rabbinical Council of America notes that significant Jewish authorities have maintained that evolutionary theory, properly understood, is not incompatible with belief in a Divine Creator, nor with the first 2 chapters of Genesis. [4]
  34. The High Council of B'nei Noah a body of non-Jews guided by the Beit Din of B'nei Noah a sub-court of the developing Sanhedrin: Science and Religion: A proper perspective through an understanding of Hebrew sources
  35. Aish HaTorah According to a possible reading of ancient commentators' description of God and nature, the world may be simultaneously young and old.
  36. On the history of eugenics and evolution, see Daniel Kevles, In the Name of Eugenics: Genetics and the Uses of Human Heredity (New York: Knopf, 1985).