Darwinism and World War One – creation.com

by Lita Cosner

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The evolutionary ideas of Charles Darwin (18091882) destructively influenced many of the Western worlds leaders in the early 20th century. In particular, intellectuals in Germany were among the earliest to embrace Darwinism enthusiastically, and to apply its concept of the survival of the fittest to human society. That is, they applied the subtitle of On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (1859). The most infamous result of this was the Holocaust,1 but social Darwinism was also a major influence in the events leading up to World War One.

In the decades leading up to World War One, intellectuals embraced Darwinism and its ethical implications as a welcome alternative to Christian belief and ethics. Ernst Haeckel (18341919), the most famous German Darwinist of the time, and notorious forger of embryo diagrams,2 believed that evolution would bring forth a complete revolution in the entire world view of humanity.3 He argued that Darwinism required the abandonment of Christian morals.

Until the advent of Darwinism, the sanctity of human life was taken for granted in European law and thought. But many German intellectuals began to argue that some had a greater right to life than others, namely, those who are deemed more valuable to society. This inequality was mainly based on race, but the Darwinists argued that there were inferior individuals within a race as well. For instance, zoologist and politician Karl Vogt (18171895) argued that a mentally handicapped child was closer in value to an ape than to his own parents.4 It should thus not be surprising that the worlds first eugenics5 society was founded in Germany, promoting the concept founded by Darwins cousin Francis Galton (18221911).6

While the belief in German superiority led the Nazis to exterminate undesirable individuals, during WW1 German Darwinists used the same idea to justify war on states which they deemed inferior.

German social Darwinists were enamoured with the vision of the master race, which in their mind was the Nordic or Germanic race. Eugenicist Alfred Ploetz (18601940) coined the term racial hygiene (Rassenhygiene), and later welcomed the Nazis as the ones who would put this into practice. While the belief in German superiority led the Nazis to exterminate undesirable individuals, during WW1 German Darwinists used the same idea to justify war on states which they deemed inferior. They believed that the destiny of the master race was to dominate or eliminate inferior races, and the most obvious way to accomplish this was through war. In their view, all races and states were in competition for survival, and those who would not wage war would perish.7 In other words war is inevitable and peace is merely an armistice in the continuous battle between races and groups for survival.8

The concept of Lebensraum or living space was one of the justifications for the Germans in both World Wars to take over their neighbouring countries. While Germany was not actually overcrowded, they believed that since one side or the other was always advancing, without war, inferior or decaying races would easily choke the growth of healthy budding elements, and a universal decadence would follow.9 Max Weber emphasized this racial competition:

Because of the view of war as an evolutionary instrument, the German leaders regarded war as a desirable option, even though they could not be sure of a victory.

Interestingly, German Darwinists were divided about whether war was beneficial for the master race. Some were pacifists, including Haeckel, because they believed that war would kill off mainly the best members of the master race, but anthropologist Otto Ammon (18421916) believed that war was the only way to test which nation was stronger and to grant victory to the fittest opponent.11 Of course, there was the problem of undesirable individuals in the German population as well; Ploetz suggested sending them to the front lines so they would be killed before those who were deemed to be more fit.12

Because of this view of war as an evolutionary instrument, the German leaders regarded war as a desirable option, even though they could not be sure of a victory. There was also a fatalistic element; they believed that it was their destiny and that they were fulfilling their preordained role in the development of the world.13

Some argue that because Darwin did not directly apply the principles of social Darwinism, the term, and its connection to evolutionary thought, is invalid. But Darwin himself said that killing in the animal kingdom was a way for evolution to progress:

Darwin simply was reluctant to apply this principle to humans, but the German social Darwinists did not share his disinclination. While there were other factors that caused World War 1, the German leaderships universal belief in social Darwinism and its anti-Christian ethical system justified their militarism and made it into a moral good.

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Social Darwinism – Dr. Hartnell’s Nutty the A.D.D. Squirrel

Eugenics is a belief and practice of improving the genetic quality of the human population. (Wow! Every word in that sentence is a landmine of problems!) Although the idea of eugenics (but not the term) is contained in the Greek philosopher Plato's Republic, the modern concept became prominent during the second half of the 19th Century. Its predecessor was the group of sociological theories known as Social Darwinism. The favorite phrases of Social Darwinism "struggle for existence" and "survival of the fittest" - when applied to humans in society, suggested that the rich were better endowed than the poor and hence more successful in life. The continual and natural sorting out of "better" and "worse" elements would, in theory, lead to continued improvement of the human species. Eugenics differs from Social Darwinism because Social Darwinism was characterized by its laissez-faire attitude, that is, letting nature take its course so that the worst elements of society were eventually be eliminated. Eugenics, on the other hand, preaches that proper breeding is the key to bettering society. In other words, you push for the higher reproduction of people with desired traits (called "positive eugenics") and prevent the reproduction of those with less-desired traits (called "negative eugenics").

This can't be serious, can it?

Oh, it is. The modern eugenicsmovement was closely associated with a sense of white Anglo-Saxon superiority. Sir Francis Galton (Charles Darwin's cousin), the founder of the English eugenics movement, for example, had been drawn to the study of human heredity and eugenics by his curiosity about what he called the hereditary "genius" in his own family... as in, "Damn, we're a smart bunch. We should be breeding more." (I'm guessing his knighthood went to his head.) The publication of Darwin's The Origin of Species in 1859 was an event that changed Galton's life. He became obsessed with the first chapter about the breeding of domestic animals (um, what?) and devoted the rest of his life to exploring the variations found in the human population. In so doing, he established a research program which categorized multiple aspects of humans, from mental characteristics to height; from facial images to fingerprint patterns. In 1883, a year after Darwin died, Galton took the Greek root words for "good" and "origin" and named his research "eugenics".

As a social movement, eugenics reached its greatest popularity in the early decades of the 20th Century. By this point, eugenics was practiced around the world and promoted by imperialist governments. Many countries enacted eugenic policies, including genetic screening, birth control, marriage restrictions, segregation (both racial segregation and segregation of the mentally ill from the rest of the population), forced sterilization, forced abortions, forced pregnancies... and genocide. The movement was especially strong in England, the U.S., and Germany from 1910-1940.

In the U.S., the eugenics movement received extensive funding from major corporations, including the Carnegie Institution and the Rockefeller Foundation. Even the inventor of corn flakes, J. H. Kellogg, launched the Race Betterment Foundation in Battle Creek, Michigan. (Yea, now your cereal tastes like shame! But trust me, that was one weird dude. Real the caption under his picture. And then go eat some cereal...) Renowned biologist, Charles B. Davenport, organized the Eugenics Record Office in New York and the American Breeder's Association (ABA). He formed the ABA specifically to "investigate and report on heredity in the human race, and to emphasize the value of superior blood and the menace to society of inferior blood." One of the ABA's biggest members wasAlexander Graham Bell. (Yea, now your phone sounds racist.)

Eugenics exerted considerable influence on popular opinion and was reflected in some state and Federal legislation. Starting with Indiana in 1907,31 states passed sterilization laws aimed at breeding out various social "misfits": the mentally retarded, criminals, and the insane. (Indiana was the first, California conducted the most, and North Carolina was the most aggressive: an IQ lower than 70 meant sterilization was appropriate.) Laws were also passed restricting marriages between members of various racial groups. Even the National Federation of Women's Clubs, the Women's Christian Temperance Union, and the National League of Women Voters were among major feminist organizations that lobbied in favor of eugenics, specifically birth control and sterilization in order to prevent unwanted children from being born into poverty and to curb passing on mental diseases or birth defects. However, Margaret Sanger, the founder of Planned Parenthood, made it abundantly clear that it wasa woman's decision and not the state whether or not to have a child. (Some see this as the start of the Pro-Choice Movement.)

The key triumph of the U.S. eugenics movement came in 1924, when a coalition of eugenicists and some big-business interests pushed through the Immigration Act of 1924, also known as the National Origins Act. The actseverely limited immigration into the U.S. from Eastern European and Mediterranean countries. Eugenicists claimed that these immigrants were inferior to Anglo-Saxons (whites) and were "polluting" the "pure" American bloodstream. By 1928, there were 376 courses in some of America's leading universities, enrolling more than 20,000 students.

After the eugenics movement was firmly in place in the U.S., it spread to Germany. Eugenicists in California began producing literatureto promoteeugenics and sterilization and sent it to German scientists and medical "professionals". By 1933, California had subjected more people to forced sterilization than all other U.S states combined.(Most werepoor, sick, and non-white.)The sterilization program engineered by the Nazis wasinspired by California's.

But wait... it gets worse.

The Rockefeller Foundation helped develop and fund various German eugenics programs, including the one that Josef Mengele worked in before he went to the concentration camp Auschwitz. (Mengele was the notorious Nazi doctor that performed horrific experiments on people.) Upon returning from Germany in 1934, where more than 5,000 people per month were being forcibly sterilized, the California eugenics leader C. M. Goethe bragged to a colleague: "You will be interested to know that your work has played a powerful part in shaping the opinions of the group of intellectuals who are behind Hitler in this epoch-making program. Everywhere I sensed that their opinions have been tremendously stimulated by American thought. I want you, my dear friend, to carry this thought with you for the rest of your life, that you have really jolted into action a great government of 60 million people."

After World War II, however, historians began to portray U.S. eugenics as different from Nazi eugenics. (Of course they did!) The movement in the U.S. was largely discredited by the fact that eugenics was central to both the theory and practice of Nazism. Still, California continued forced sterilizations on prisoners as late as the mid-1960s, mostly because California's long-time attorney general was a big supporter of the practice, and it wasn't formally outlawed there until 1979. (In fact, from 2006-2010, 148 women were illegally sterilized in California's prisons.)

Eugenicsis rightfully andseverely criticized for what can be calledovert racial bias, subjectivity in the use of evidence, and lack of scientific "proof".Eugenics isa shameful reminder of what happens when science mixes with racism. But don't worry. You can sleep on this fun fact: eugenics is still officially permitted in the U.S. today. WHAT?!

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Social Darwinism - Dr. Hartnell's Nutty the A.D.D. Squirrel

Social Darwinism – University of Colorado Boulder

Social Darwinism

I. Introduction

Social Darwinism, term coined in the late 19th century to describe the idea that humans, like animals and plants, compete in a struggle for existence in which natural selection results in "survival of the fittest." Social Darwinists base their beliefs on theories of evolution developed by British naturalist Charles Darwin. Some social Darwinists argue that governments should not interfere with human competition by attempting to regulate the economy or cure social ills such as poverty. Instead, they advocate a laissez-faire political and economic system that favors competition and self-interest in social and business affairs. Social Darwinists typically deny that they advocate a "law of the jungle." But most propose arguments that justify imbalances of power between individuals, races, and nations because they consider some people more fit to survive than others.

The term social Darwinist is applied loosely to anyone who interprets human society primarily in terms of biology, struggle, competition, or natural law (a philosophy based on what are considered the permanent characteristics of human nature). Social Darwinism characterizes a variety of past and present social policies and theories, from attempts to reduce the power of government to theories exploring the biological causes of human behavior. Many people believe that the concept of social Darwinism explains the philosophical rationalization behind racism, imperialism, and capitalism. The term has negative implications for most people because they consider it a rejection of compassion and social responsibility.

II. Origins

Social Darwinism originated in Britain during the second half of the 19th century. Darwin did not address human evolution in his most famous study, On the Origin of Species (1859), which focused on the evolution of plants and animals. He applied his theories of natural selection specifically to people in The Descent of Man (1871), a work that critics interpreted as justifying cruel social policies at home and imperialism abroad. The Englishman most associated with early social Darwinism, however, was sociologist Herbert Spencer. Spencer coined the phrase "survival of the fittest" to describe the outcome of competition between social groups. In Social Statics (1850) and other works, Spencer argued that through competition social evolution would automatically produce prosperity and personal liberty unparalleled in human history.

In the United States, Spencer gained considerable support among intellectuals and some businessmen, including steel manufacturer Andrew Carnegie, who served as Spencer's host during his visit to the United States in 1883. The most prominent American social Darwinist of the 1880s was William Graham Sumner, who on several occasions told audiences that there was no alternative to the "survival of the fittest" theory. Critics of social Darwinism seized on these comments to argue that Sumner advocated a "dog-eat-dog" philosophy of human behavior that justified oppressive social policies. Some later historians have argued that Sumner's critics took his statements out of context and misrepresented his views.

III. Hereditarianism

Studies of heredity contributed another variety of social Darwinism in the late 19th century. In Hereditary Genius (1869), Sir Francis Galton, a British scientist and Darwin's cousin, argued that biological inheritance is far more important than environment in determining character and intelligence. This theory, known as hereditarianism, met considerable resistance, especially in the United States. Sociologists and biologists who criticized hereditarianism believed that changes in the environment could produce physical changes in the individual that would be passed on to future generations, a theory proposed by French biologist Jean-Baptiste Lamarck in the early 19th century. After 1890, hereditarianism gained increasing support, due in part to the work of German biologist August Weismann. Weismann reemphasized the role of natural selection by arguing that a person's characteristics are determined genetically at conception.

IV. The Struggle School

Toward the end of the 19th century, another strain of social Darwinism was developed by supporters of the struggle school of sociology. English journalist Walter Bagehot expressed the fundamental ideas of the struggle school in Physics and Politics (1872), a book that describes the historical evolution of social groups into nations. Bagehot argued that these nations evolved principally by succeeding in conflicts with other groups. For many political scientists, sociologists, and military strategists, this strain of social Darwinism justified overseas expansion by nations (imperialism) during the 1890s. In the United States, historian John Fiske and naval strategist Alfred Thayer Mahan drew from the principles of social Darwinism to advocate foreign expansion and the creation of a strong military.

V. Reform Darwinism

After 1890, social reformers used Darwinism to advocate a stronger role for government and the introduction of various social policies. This movement became known as reform Darwinism. Reform Darwinists argued that human beings need new ideas and institutions as they adapt to changing conditions. For example, U.S. Supreme Court Justice Oliver Wendell Holmes, Jr. reasoned that the Constitution of the United States should be reinterpreted in light of changing circumstances in American society.

Some reformers used the principles of evolution to justify sexist and racist ideas that undercut their professed belief in equality. For example, the most extreme type of reform Darwinism was eugenics, a term coined by Sir Francis Galton in 1883 from the Greek word egenv, meaning well-born. Eugenists claimed that particular racial or social groupsusually wealthy Anglo-Saxonswere "naturally" superior to other groups. They proposed to control human heredity by passing laws that forbid marriage between races or that restrict breeding for various social "misfits" such as criminals or the mentally ill.

VI. Social Darwinism in the 20th Century

Although social Darwinism was highly influential at the beginning of the 20th century, it rapidly lost popularity and support after World War I (1914-1918). During the 1920s and 1930s many political observers blamed it for contributing to German militarism and the rise of Nazism (see National Socialism). During this same period, advances in anthropology also discredited social Darwinism. German American anthropologist Franz Boas and American anthropologists Margaret Mead and Ruth Benedict showed that human culture sets people apart from animals. By shifting the emphasis away from biology and onto culture, these anthropologists undermined social Darwinism's biological foundations. Eugenics was discredited by a better understanding of genetics and eventually disgraced by Nazi dictator Adolf Hitler's use of eugenic arguments to create a "master race." During World War II (1939-1945), the Nazis killed several million Jews, Roma (Gypsies), and members of other groups, believing them inferior to an idealized Aryan race.

Social theories based on biology gained renewed support after 1953, when American biologist James Watson and British biologist Francis Crick successfully described the structure of the DNA molecule, the building block of all life. During the 1960s anthropologists interested in the influence of DNA on human behavior produced studies of the biological basis of aggression, territoriality, mate selection, and other behavior common to people and animals. Books on this theme, such as Desmond Morris's Naked Ape (1967) and Lionel Tiger's Men in Groups (1969), became best-sellers. In the early 1970s American psychologist Richard J. Herrnstein revived the social Darwinist argument that intelligence is mostly determined by biology rather than by environmental influences.

During the 1960s, British biologist W. D. Hamilton and American biologist Robert L. Trivers produced separate studies showing that the self-sacrificing behavior of some members of a group serves the genetic well-being of the group as a whole. American biologist Edward O. Wilson drew on these theories in Sociobiology: the New Synthesis (1975), where he argued that genetics exerts a greater influence on human behavior than scientists had previously believed. Wilson claimed that human behavior cannot be understood without taking both biology and culture into account. Wilson's views became the foundations of a new sciencesociobiologyand were later popularized in such studies as Richard Dawkins'The Selfish Gene (1976). Wilson's critics have alleged that sociobiology is simply another version of social Darwinism. They claim that it downplays the role of culture in human societies and justifies poverty and warfare in the name of natural selection. Such criticism has led to a decline in the influence of sociobiology and other forms of social Darwinism.

Contributed By:

Robert C. Bannister, B.A., M.A., Ph.D.

Professor of History, Swarthmore College. Author of Social Darwinism: Science and Myth and On Liberty, Society, and Politics: The Essential Essays of William Graham Sumner.

HOW TO CITE THIS ARTICLE

"Social Darwinism," Microsoft Encarta Online Encyclopedia 2000

http://encarta.msn.com 1997-2000 Microsoft Corporation. All rights reserved.

1993-2000 Microsoft Corporation.

All rights reserved.

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Social Darwinism - University of Colorado Boulder

Natural selection – Wikipedia

Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in heritable traits of a population over time. Charles Darwin popularised the term "natural selection", and compared it with artificial selection.

Variation exists within all populations of organisms. This occurs partly because random mutations arise in the genome of an individual organism, and offspring can inherit such mutations. Throughout the lives of the individuals, their genomes interact with their environments to cause variations in traits. The environment of a genome includes the molecular biology in the cell, other cells, other individuals, populations, species, as well as the abiotic environment. Individuals with certain variants of the trait may survive and reproduce more than individuals with other, less successful, variants. Therefore, the population evolves. Factors that affect reproductive success are also important, an issue that Darwin developed in his ideas on sexual selection (now often included in natural selection) and fecundity selection.

Natural selection acts on the phenotype, or the observable characteristics of an organism, but the genetic (heritable) basis of any phenotype that gives a reproductive advantage may become more common in a population. Over time, this process can result in populations that specialise for particular ecological niches (microevolution) and may eventually result in the emergence of new species (macroevolution). In other words, natural selection is a key process in the evolution of a population. Natural selection can be contrasted with artificial selection, in which humans intentionally choose specific traits, whereas in natural selection there is no intentional choice.

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

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

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

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

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

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

The classical arguments were reintroduced in the 18th century by Pierre Louis Maupertuis[11] and others, including Darwin's grandfather, Erasmus Darwin.

Until the early 19th century, the prevailing view in Western societies was that differences between individuals of a species were uninteresting departures from their Platonic ideals (or typus) of created kinds. However, the theory of uniformitarianism in geology promoted the idea that simple, weak forces could act continuously over long periods of time to produce radical changes in the Earth's landscape. The success of this theory raised awareness of the vast scale of geological time and made plausible the idea that tiny, virtually imperceptible changes in successive generations could produce consequences on the scale of differences between species.[12]

The early 19th-century zoologist Jean-Baptiste Lamarck suggested the inheritance of acquired characteristics as a mechanism for evolutionary change; adaptive traits acquired by an organism during its lifetime could be inherited by that organism's progeny, eventually causing transmutation of species.[13] This theory, Lamarckism, was an influence on the Soviet biologist Trofim Lysenko's antagonism to mainstream genetic theory as late as the mid 20th century.[14]

Between 1835 and 1837, the zoologist Edward Blyth worked on the area of variation, artificial selection, and how a similar process occurs in nature. Darwin acknowledged Blyth's ideas in the first chapter on variation of On the Origin of Species.[15]

In 1859, Charles Darwin set out his theory of evolution by natural selection as an explanation for adaptation and speciation. He defined natural selection as the "principle by which each slight variation [of a trait], if useful, is preserved."[16] The concept was simple but powerful: individuals best adapted to their environments are more likely to survive and reproduce. As long as there is some variation between them and that variation is heritable, there will be an inevitable selection of individuals with the most advantageous variations. If the variations are heritable, then differential reproductive success leads to a progressive evolution of particular populations of a species, and populations that evolve to be sufficiently different eventually become different species.[17][18]

Darwin's ideas were inspired by the observations that he had made on the second voyage of HMS Beagle (18311836), and by the work of a political economist, Thomas Robert Malthus, who, in An Essay on the Principle of Population (1798), noted that population (if unchecked) increases exponentially, whereas the food supply grows only arithmetically; thus, inevitable limitations of resources would have demographic implications, leading to a "struggle for existence."[19] When Darwin read Malthus in 1838 he was already primed by his work as a naturalist to appreciate the "struggle for existence" in nature and it struck him that as population outgrew resources, "favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species."[20] Darwin wrote:

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

Once he had his theory, Darwin was meticulous about gathering and refining evidence before making his idea public. He was in the process of writing his "big book" to present his research when the naturalist Alfred Russel Wallace independently conceived of the principle and described it in an essay he sent to Darwin to forward to Charles Lyell. Lyell and Joseph Dalton Hooker decided (without Wallace's knowledge) to present his essay together with unpublished writings that Darwin had sent to fellow naturalists, and On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection was read to the Linnean Society of London announcing co-discovery of the principle in July 1858.[22] Darwin published a detailed account of his evidence and conclusions in On the Origin of Species in 1859. In the 3rd edition of 1861 Darwin acknowledged that otherslike William Charles Wells in 1813, and Patrick Matthew in 1831had proposed similar ideas, but had neither developed them nor presented them in notable scientific publications.[23]

Darwin thought of natural selection by analogy to how farmers select crops or livestock for breeding, which he called "artificial selection"; in his early manuscripts he referred to a "Nature" which would do the selection. At the time, other mechanisms of evolution such as evolution by genetic drift were not yet explicitly formulated, and Darwin believed that selection was likely only part of the story: "I am convinced that Natural Selection has been the main but not exclusive means of modification."[24] In a letter to Charles Lyell in September 1860, Darwin regretted the use of the term "Natural Selection," preferring the term "Natural Preservation."[25]

For Darwin and his contemporaries, natural selection was in essence synonymous with evolution by natural selection. After the publication of On the Origin of Species,[26] educated people generally accepted that evolution had occurred in some form. However, natural selection remained controversial as a mechanism, partly because it was perceived to be too weak to explain the range of observed characteristics of living organisms, and partly because even supporters of evolution balked at its "unguided" and non-progressive nature,[27] a response that has been characterised as the single most significant impediment to the idea's acceptance.[28] However, some thinkers enthusiastically embraced natural selection; after reading Darwin, Herbert Spencer introduced the phrase survival of the fittest, which became a popular summary of the theory.[29][30] The fifth edition of On the Origin of Species published in 1869 included Spencer's phrase as an alternative to natural selection, with credit given: "But the expression often used by Mr. Herbert Spencer of the Survival of the Fittest is more accurate, and is sometimes equally convenient."[31] Although the phrase is still often used by non-biologists, modern biologists avoid it because it is tautological if "fittest" is read to mean "functionally superior" and is applied to individuals rather than considered as an averaged quantity over populations.[32]

Natural selection relies crucially on the idea of heredity, but developed before the basic concepts of genetics. Although the Moravian monk Gregor Mendel, the father of modern genetics, was a contemporary of Darwin's, his work lay in obscurity, only being rediscovered in 1900.[33] Only after the mid-20th century integration of evolution with Mendel's laws of inheritance, the so-called modern evolutionary synthesis, did scientists generally come to accept natural selection.[34][35] The synthesis grew from advances in different fields. Ronald Fisher developed the required mathematical language and wrote The Genetical Theory of Natural Selection (1930).[36]J. B. S. Haldane introduced the concept of the "cost" of natural selection.[37][38]Sewall Wright elucidated the nature of selection and adaptation.[39]Theodosius Dobzhansky established the idea that mutation, by creating genetic diversity, supplied the raw material for natural selection, in his book Genetics and the Origin of Species (1937).[40][41]Ernst Mayr recognised the key importance of reproductive isolation for speciation in his Systematics and the Origin of Species (1942).[42]W. D. Hamilton conceived of kin selection in 1964.[43][44] This synthesis cemented natural selection as the foundation of evolutionary theory, where it remains today. A second synthesis was brought about at the end of the 20th century by molecular genetics, creating the field of evolutionary developmental biology (evo-devo), which seeks to explain evolution at a molecular level.[45][46][47]

The term natural selection is most often defined to operate on heritable traits, because these directly participate in evolution. However, natural selection is "blind" in the sense that changes in phenotype can give a reproductive advantage regardless of whether or not the trait is heritable. Following Darwin's primary usage, the term is used to refer both to the evolutionary consequence of blind selection and to its mechanisms.[26][36][48][49] It is sometimes helpful to explicitly distinguish between selection's mechanisms and its effects; when this distinction is important, scientists define "(phenotypic) natural selection" specifically as "those mechanisms that contribute to the selection of individuals that reproduce", without regard to whether the basis of the selection is heritable.[50][51][52] Traits that cause greater reproductive success of an organism are said to be selected for, while those that reduce success are selected against.[53]

Natural variation occurs among the individuals of any population of organisms. Some differences may improve an individual's chances of surviving and reproducing, so that its lifetime reproductive rate is increased: it leaves more offspring. If the traits that give these individuals a reproductive advantage are also heritable, that is, passed from parent to offspring, then there will be differential reproduction, that is, a slightly higher proportion of fast rabbits or efficient algae in the next generation. Even if the reproductive advantage is very slight, over many generations any advantageous heritable trait becomes dominant in the population. In this way the natural environment of an organism "selects for" traits that confer a reproductive advantage, causing evolutionary change, as Darwin described.[54] This gives the appearance of purpose, but in natural selection there is no intentional choice. Artificial selection is purposive where natural selection is not, though biologists often use teleological language to describe it.[55]

The peppered moth exists in both light and dark colours in Great Britain, but during the industrial revolution, many of the trees on which the moths rested became blackened by soot, giving the dark-coloured moths an advantage in hiding from predators. This gave dark-coloured moths a better chance of surviving to produce dark-coloured offspring, and in just fifty years from the first dark moth being caught, nearly all of the moths in industrial Manchester were dark. The balance was reversed by the effect of the Clean Air Act 1956, and the dark moths became rare again, demonstrating the influence of natural selection on peppered moth evolution.[56]

The concept of fitness is central to natural selection. In broad terms, individuals that are more "fit" have better potential for survival, as in the well-known phrase "survival of the fittest", but the precise meaning of the term is much more subtle. Modern evolutionary theory defines fitness not by how long an organism lives, but by how successful it is at reproducing. If an organism lives half as long as others of its species, but has twice as many offspring surviving to adulthood, its genes become more common in the adult population of the next generation. Though natural selection acts on individuals, the effects of chance mean that fitness can only really be defined "on average" for the individuals within a population. The fitness of a particular genotype corresponds to the average effect on all individuals with that genotype.[57]

In biology, competition is an interaction between organisms in which the fitness of one is lowered by the presence of another. This may be because both rely on a limited supply of a resource such as food, water, or territory.[58] Competition may be within or between species, and may be direct or indirect.[59] Species less suited to compete should in theory either adapt or die out, since competition plays a powerful role in natural selection, but according to the "room to roam" theory it may be less important than expansion among larger clades.[59][60]

Competition is modelled by r/K selection theory, which is based on Robert MacArthur and E. O. Wilson's work on island biogeography.[61] In this theory, selective pressures drive evolution in one of two stereotyped directions: r- or K-selection.[62] These terms, r and K, can be illustrated in a logistic model of population dynamics:[63]

where r is the growth rate of the population (N), and K is the carrying capacity of its local environmental setting. Typically, r-selected species exploit empty niches, and produce many offspring, each with a relatively low probability of surviving to adulthood. In contrast, K-selected species are strong competitors in crowded niches, and invest more heavily in much fewer offspring, each with a relatively high probability of surviving to adulthood.[63]

Natural selection can act on any heritable phenotypic trait,[64] and selective pressure can be produced by any aspect of the environment, including sexual selection and competition with members of the same or other species.[65][66] However, this does not imply that natural selection is always directional and results in adaptive evolution; natural selection often results in the maintenance of the status quo by eliminating less fit variants.[54]

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

Selection has different effects on traits. Stabilizing selection acts to hold a trait at a stable optimum, and in the simplest case all deviations from this optimum are selectively disadvantageous. Directional selection favours extreme values of a trait. The uncommon disruptive selection also acts during transition periods when the current mode is sub-optimal, but alters the trait in more than one direction. In particular, if the trait is quantitative and univariate then both higher and lower trait levels are favoured. Disruptive selection can be a precursor to speciation.[54]

Alternatively, selection can be divided according to its effect on genetic diversity. Purifying or negative selection acts to remove genetic variation from the population (and is opposed by de novo mutation, which introduces new variation.[67][68] In contrast, balancing selection acts to maintain genetic variation in a population, even in the absence of de novo mutation, by negative frequency-dependent selection. One mechanism for this is heterozygote advantage, where individuals with two different alleles have a selective advantage over individuals with just one allele. The polymorphism at the human ABO blood group locus has been explained in this way.[69]

Another option is to classify selection by the life cycle stage at which it acts. Some biologists recognise just two types: viability (or survival) selection, which acts to increase an organism's probability of survival, and fecundity (or fertility or reproductive) selection, which acts to increase the rate of reproduction, given survival. Others split the life cycle into further components of selection. Thus viability and survival selection may be defined separately and respectively as acting to improve the probability of survival before and after reproductive age is reached, while fecundity selection may be split into additional sub-components including sexual selection, gametic selection, acting on gamete survival, and compatibility selection, acting on zygote formation.[70]

Selection can also be classified by the level or unit of selection. Individual selection acts on the individual, in the sense that adaptations are "for" the benefit of the individual, and result from selection among individuals. Gene selection acts directly at the level of the gene. In kin selection and intragenomic conflict, gene-level selection provides a more apt explanation of the underlying process. Group selection, if it occurs, acts on groups of organisms, on the assumption that groups replicate and mutate in an analogous way to genes and individuals. There is an ongoing debate over the degree to which group selection occurs in nature.[71]

Finally, selection can be classified according to the resource being competed for. Sexual selection results from competition for mates. Sexual selection typically proceeds via fecundity selection, sometimes at the expense of viability. Ecological selection is natural selection via any means other than sexual selection, such as kin selection, competition, and infanticide. Following Darwin, natural selection is sometimes defined as ecological selection, in which case sexual selection is considered a separate mechanism.[72]

Sexual selection as first articulated by Darwin and Wallace[73] refers specifically to competition for mates,[74] which can be intrasexual, between individuals of the same sex, that is malemale competition, or intersexual, where one gender chooses mates, most often with males displaying and females choosing.[75] However, in some species, mate choice is primarily by males, as in some fishes of the family Syngnathidae.[76][77]

Phenotypic traits can be displayed in one sex and desired in the other sex, causing a positive feedback loop called a Fisherian runaway, for example, the extravagant plumage of some male birds. An alternate theory proposed by the same Ronald Fisher in 1930 is the sexy son hypothesis, that mothers want promiscuous sons to give them large numbers of grandchildren and so choose promiscuous fathers for their children. Aggression between members of the same sex is sometimes associated with very distinctive features, such as the antlers of stags, which are used in combat with other stags. More generally, intrasexual selection is often associated with sexual dimorphism, including differences in body size between males and females of a species.[75]

Natural selection is seen in action in the development of antibiotic resistance in microorganisms. Since the discovery of penicillin in 1928, antibiotics have been used to fight bacterial diseases. The widespread misuse of antibiotics has selected for microbial resistance to antibiotics in clinical use, to the point that the methicillin-resistant Staphylococcus aureus (MRSA) has been described as a "superbug" because of the threat it poses to health and its relative invulnerability to existing drugs.[78] Response strategies typically include the use of different, stronger antibiotics; however, new strains of MRSA have recently emerged that are resistant even to these drugs.[79] This is an evolutionary arms race, in which bacteria develop strains less susceptible to antibiotics, while medical researchers attempt to develop new antibiotics that can kill them. A similar situation occurs with pesticide resistance in plants and insects. Arms races are not necessarily induced by man; a well-documented example involves the spread of a gene in the butterfly Hypolimnas bolina suppressing male-killing activity by Wolbachia bacteria parasites on the island of Samoa, where the spread of the gene is known to have occurred over a period of just five years [80][81]

A prerequisite for natural selection to result in adaptive evolution, novel traits and speciation is the presence of heritable genetic variation that results in fitness differences. Genetic variation is the result of mutations, genetic recombinations and alterations in the karyotype (the number, shape, size and internal arrangement of the chromosomes). Any of these changes might have an effect that is highly advantageous or highly disadvantageous, but large effects are rare. In the past, most changes in the genetic material were considered neutral or close to neutral because they occurred in noncoding DNA or resulted in a synonymous substitution. However, many mutations in non-coding DNA have deleterious effects.[82][83] Although both mutation rates and average fitness effects of mutations are dependent on the organism, a majority of mutations in humans are slightly deleterious.[84]

Some mutations occur in "toolkit" or regulatory genes. Changes in these often have large effects on the phenotype of the individual because they regulate the function of many other genes. Most, but not all, mutations in regulatory genes result in non-viable embryos. Some nonlethal regulatory mutations occur in HOX genes in humans, which can result in a cervical rib[85] or polydactyly, an increase in the number of fingers or toes.[86] When such mutations result in a higher fitness, natural selection favours these phenotypes and the novel trait spreads in the population. Established traits are not immutable; traits that have high fitness in one environmental context may be much less fit if environmental conditions change. In the absence of natural selection to preserve such a trait, it becomes more variable and deteriorate over time, possibly resulting in a vestigial manifestation of the trait, also called evolutionary baggage. In many circumstances, the apparently vestigial structure may retain a limited functionality, or may be co-opted for other advantageous traits in a phenomenon known as preadaptation. A famous example of a vestigial structure, the eye of the blind mole-rat, is believed to retain function in photoperiod perception.[87]

Speciation requires a degree of reproductive isolationthat is, a reduction in gene flow. However, it is intrinsic to the concept of a species that hybrids are selected against, opposing the evolution of reproductive isolation, a problem that was recognised by Darwin. The problem does not occur in allopatric speciation with geographically separated populations, which can diverge with different sets of mutations. E. B. Poulton realized in 1903 that reproductive isolation could evolve through divergence, if each lineage acquired a different, incompatible allele of the same gene. Selection against the heterozygote would then directly create reproductive isolation, leading to the BatesonDobzhanskyMuller model, further elaborated by H. Allen Orr and Michael Turelli.[88]

Natural selection acts on an organism's phenotype, or physical characteristics. Phenotype is determined by an organism's genetic make-up (genotype) and the environment in which the organism lives. When different organisms in a population possess different versions of a gene for a certain trait, each of these versions is known as an allele. It is this genetic variation that underlies differences in phenotype. An example is the ABO blood type antigens in humans, where three alleles govern the phenotype.[89]

Some traits are governed by only a single gene, but most traits are influenced by the interactions of many genes. A variation in one of the many genes that contributes to a trait may have only a small effect on the phenotype; together, these genes can produce a continuum of possible phenotypic values.[90]

When some component of a trait is heritable, selection alters the frequencies of the different alleles, or variants of the gene that produces the variants of the trait. Selection can be divided into three classes, on the basis of its effect on allele frequencies: directional, stabilizing, and purifying selection.[91] Directional selection occurs when an allele has a greater fitness than others, so that it increases in frequency, gaining an increasing share in the population. This process can continue until the allele is fixed and the entire population shares the fitter phenotype.[92] Far more common is stabilizing selection, which lowers the frequency of alleles that have a deleterious effect on the phenotype that is, produce organisms of lower fitness. This process can continue until the allele is eliminated from the population. Purifying selection conserves functional genetic features, such as protein-coding genes or regulatory sequences, over time by selective pressure against deleterious variants.[93]

Some forms of balancing selection do not result in fixation, but maintain an allele at intermediate frequencies in a population. This can occur in diploid species (with pairs of chromosomes) when heterozygous individuals (with just one copy of the allele) have a higher fitness than homozygous individuals (with two copies). This is called heterozygote advantage or over-dominance, of which the best-known example is the resistance to malaria in humans heterozygous for sickle-cell anaemia. Maintenance of allelic variation can also occur through disruptive or diversifying selection, which favours genotypes that depart from the average in either direction (that is, the opposite of over-dominance), and can result in a bimodal distribution of trait values. Finally, balancing selection can occur through frequency-dependent selection, where the fitness of one particular phenotype depends on the distribution of other phenotypes in the population. The principles of game theory have been applied to understand the fitness distributions in these situations, particularly in the study of kin selection and the evolution of reciprocal altruism.[94][95]

A portion of all genetic variation is functionally neutral, producing no phenotypic effect or significant difference in fitness. Motoo Kimura's neutral theory of molecular evolution by genetic drift proposes that this variation accounts for a large fraction of observed genetic diversity.[96] Neutral events can radically reduce genetic variation through population bottlenecks.[97] which among other things can cause the founder effect in initially small new populations.[98] When genetic variation does not result in differences in fitness, selection cannot directly affect the frequency of such variation. As a result, the genetic variation at those sites is higher than at sites where variation does influence fitness.[91] However, after a period with no new mutations, the genetic variation at these sites is eliminated due to genetic drift. Natural selection reduces genetic variation by eliminating maladapted individuals, and consequently the mutations that caused the maladaptation. At the same time, new mutations occur, resulting in a mutationselection balance. The exact outcome of the two processes depends both on the rate at which new mutations occur and on the strength of the natural selection, which is a function of how unfavourable the mutation proves to be.[99]

Genetic linkage occurs when the loci of two alleles are in close proximity on a chromosome. During the formation of gametes, recombination reshuffles the alleles. The chance that such a reshuffle occurs between two alleles is inversely related to the distance between them. Selective sweeps occur when an allele becomes more common in a population as a result of positive selection. As the prevalence of one allele increases, closely linked alleles can also become more common by "genetic hitchhiking", whether they are neutral or even slightly deleterious. A strong selective sweep results in a region of the genome where the positively selected haplotype (the allele and its neighbours) are in essence the only ones that exist in the population. Selective sweeps can be detected by measuring linkage disequilibrium, or whether a given haplotype is overrepresented in the population. Since a selective sweep also results in selection of neighbouring alleles, the presence of a block of strong linkage disequilibrium might indicate a 'recent' selective sweep near the centre of the block.[100]

Background selection is the opposite of a selective sweep. If a specific site experiences strong and persistent purifying selection, linked variation tends to be weeded out along with it, producing a region in the genome of low overall variability. Because background selection is a result of deleterious new mutations, which can occur randomly in any haplotype, it does not produce clear blocks of linkage disequilibrium, although with low recombination it can still lead to slightly negative linkage disequilibrium overall.[101]

Darwin's ideas, along with those of Adam Smith and Karl Marx, had a profound influence on 19th century thought, including his radical claim that "elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner" evolved from the simplest forms of life by a few simple principles.[102] This inspired some of Darwin's most ardent supportersand provoked the strongest opposition. Natural selection had the power, according to Stephen Jay Gould, to "dethrone some of the deepest and most traditional comforts of Western thought", such as the belief that humans have a special place in the world.[103]

In the words of the philosopher Daniel Dennett, "Darwin's dangerous idea" of evolution by natural selection is a "universal acid," which cannot be kept restricted to any vessel or container, as it soon leaks out, working its way into ever-wider surroundings.[104] Thus, in the last decades, the concept of natural selection has spread from evolutionary biology to other disciplines, including evolutionary computation, quantum Darwinism, evolutionary economics, evolutionary epistemology, evolutionary psychology, and cosmological natural selection. This unlimited applicability has been called universal Darwinism.[105]

How life originated from inorganic matter remains an unresolved problem in biology. One prominent hypothesis is that life first appeared in the form of short self-replicating RNA polymers.[106] On this view, life may have come into existence when RNA chains first experienced the basic conditions, as conceived by Charles Darwin, for natural selection to operate. These conditions are: heritability, variation of type, and competition for limited resources. The fitness of an early RNA replicator would likely have been a function of adaptive capacities that were intrinsic (i.e., determined by the nucleotide sequence) and the availability of resources.[107][108] The three primary adaptive capacities could logically have been: (1) the capacity to replicate with moderate fidelity (giving rise to both heritability and variation of type), (2) the capacity to avoid decay, and (3) the capacity to acquire and process resources.[107][108] These capacities would have been determined initially by the folded configurations (including those configurations with ribozyme activity) of the RNA replicators that, in turn, would have been encoded in their individual nucleotide sequences.[109]

In 1881, the embryologist Wilhelm Roux published Der Kampf der Theile im Organismus (The Struggle of Parts in the Organism) in which he suggested that the development of an organism results from a Darwinian competition between the parts of the embryo, occurring at all levels, from molecules to organs.[110] In recent years, a modern version of this theory has been proposed by Jean-Jacques Kupiec. According to this cellular Darwinism, random variation at the molecular level generates diversity in cell types whereas cell interactions impose a characteristic order on the developing embryo.[111]

The social implications of the theory of evolution by natural selection also became the source of continuing controversy. Friedrich Engels, a German political philosopher and co-originator of the ideology of communism, wrote in 1872 that "Darwin did not know what a bitter satire he wrote on mankind, and especially on his countrymen, when he showed that free competition, the struggle for existence, which the economists celebrate as the highest historical achievement, is the normal state of the animal kingdom."[112] Herbert Spencer and the eugenics advocate Francis Galton's interpretation of natural selection as necessarily progressive, leading to supposed advances in intelligence and civilisation, became a justification for colonialism, eugenics, and social Darwinism. For example, in 1940, Konrad Lorenz, in writings that he subsequently disowned, used the theory as a justification for policies of the Nazi state. He wrote "...selection for toughness, heroism, and social utility... must be accomplished by some human institution, if mankind, in default of selective factors, is not to be ruined by domestication-induced degeneracy. The racial idea as the basis of our state has already accomplished much in this respect."[113] Others have developed ideas that human societies and culture evolve by mechanisms analogous to those that apply to evolution of species.[114]

More recently, work among anthropologists and psychologists has led to the development of sociobiology and later of evolutionary psychology, a field that attempts to explain features of human psychology in terms of adaptation to the ancestral environment. The most prominent example of evolutionary psychology, notably advanced in the early work of Noam Chomsky and later by Steven Pinker, is the hypothesis that the human brain has adapted to acquire the grammatical rules of natural language.[115] Other aspects of human behaviour and social structures, from specific cultural norms such as incest avoidance to broader patterns such as gender roles, have been hypothesised to have similar origins as adaptations to the early environment in which modern humans evolved. By analogy to the action of natural selection on genes, the concept of memes"units of cultural transmission," or culture's equivalents of genes undergoing selection and recombinationhas arisen, first described in this form by Richard Dawkins in 1976[116] and subsequently expanded upon by philosophers such as Daniel Dennett as explanations for complex cultural activities, including human consciousness.[117]

In 1922, Alfred J. Lotka proposed that natural selection might be understood as a physical principle that could be described in terms of the use of energy by a system,[118][119] a concept later developed by Howard T. Odum as the maximum power principle in thermodynamics, whereby evolutionary systems with selective advantage maximise the rate of useful energy transformation.[120]

The principles of natural selection have inspired a variety of computational techniques, such as "soft" artificial life, that simulate selective processes and can be highly efficient in 'adapting' entities to an environment defined by a specified fitness function.[121] For example, a class of heuristic optimisation algorithms known as genetic algorithms, pioneered by John Henry Holland in the 1970s and expanded upon by David E. Goldberg,[122] identify optimal solutions by simulated reproduction and mutation of a population of solutions defined by an initial probability distribution.[123] Such algorithms are particularly useful when applied to problems whose energy landscape is very rough or has many local minima.[124]

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Natural selection - Wikipedia

Difference between Darwinism and Neo-Darwinism | Major …

The evolutionary idea contributed proposed by Charles Darwin called Darwinism or Natural selection theory, explaining the mechanism of evolution is clearly stated in his book Origin of species.

The important postulates of the theory are: Over production, Struggle for existence, Variations, Survival of the fittest, and Origin of species.

A few Neo Darwinism Supporters are Romanes, Wallace, Fisher, Huxley, Ford, Haldane, Goldschmidt, Sewall Wright, Ernst Haeckel, August Weismann, Mendel, Dobazhansky , Kettlewell and Herbert Spencer.

The Neo Darwinism has the following ideas: Experimental evidences and Answers to the objections

Darwinism vsNeo-Darwinism

2. It considers all inheritable favourable variation

3. It does not explain the reason for variation

4. In Darwins theory, the basic unit of evolution is an individual

5. It does not consider reproductive isolation as a major factor in new species formation

6. In Darwins theory, natural selection is the survival of the fittest and removal of the unfit ones during the course of time

Neo-Darwinism

1. It is the modification of original concept postulated by Darwin and Wallace based on data obtained from genetic research

2. It considers only inheritable genetic variation (mutations) for evolution

3. It explains the reason for variations

4. In Neo-Darwinism, the basic unit of evolution is a population

5. It consider reproductive isolation as a major essential factor in speciation

6. In Neo-Darwinism, natural selection refers to the differential reproduction leading to the changes in gene frequency

The rest is here:

Difference between Darwinism and Neo-Darwinism | Major ...

Why Darwinism Is False | Center for Science and Culture

Jonathan Wells Discovery Institute May 18, 2009 Print Article

Jerry A. Coyne is a professor in the Department of Ecology and Evolution at The University of Chicago. In Why Evolution is True, he summarizes Darwinismthe modern theory of evolutionas follows: Life on earth evolved gradually beginning with one primitive speciesperhaps a self-replicating moleculethat lived more than 3.5 billion years ago; it then branched out over time, throwing off many new and diverse species; and the mechanism for most (but not all) of evolutionary change is natural selection.1

Coyne further explains that evolution simply means that a species undergoes genetic change over time. That is, over many generations a species can evolve into something quite different, and those differences are based on changes in the DNA, which originate as mutations. The species of animals and plants living today werent around in the past, but are descended from those that lived earlier.2

According to Coyne, however, if evolution meant only gradual genetic change within a species, wed have only one species todaya single highly evolved descendant of the first species. Yet we have many How does this diversity arise from one ancestral form? It arises because of splitting, or, more accurately, speciation, which simply means the evolution of different groups that cant interbreed.3

If Darwinian theory were true, we should be able to find some cases of speciation in the fossil record, with one line of descent dividing into two or more. And we should be able to find new species forming in the wild. Furthermore, we should be able to find examples of species that link together major groups suspected to have common ancestry, like birds with reptiles and fish with amphibians. Finally, there are facts that make sense only in light of the theory of evolution but do not make sense in the light of creation or design. These include patterns of species distribution on the earths surface, peculiarities of how organisms develop from embryos, and the existence of vestigial features that are of no apparent use. Coyne concludes his introduction with the bold statement that all the evidenceboth old and newleads ineluctably to the conclusion that evolution is true.4

Of course, evolution is undeniably true if it means simply that existing species can change in minor ways over time, or that many species living today did not exist in the past. But Darwins claim that all species are modified descendants of a common ancestor, and Coynes claim that DNA mutations and natural selection have produced those modifications, are not so undeniably true. Coyne devotes the remainder of his book to providing evidence for them.

Fossils

Coyne turns first to the fossil record. We should be able, he writes, to find some evidence for evolutionary change in the fossil record. The deepest (and oldest) layers of rock would contain the fossils of more primitive species, and some fossils should become more complex as the layers of rock become younger, with organisms resembling present-day species found in the most recent layers. And we should be able to see some species changing over time, forming lineages showing descent with modification (adaptation). In particular, later species should have traits that make them look like the descendants of earlier ones.5

In The Origin of Species, Charles Darwin acknowledged that the fossil record presented difficulties for his theory. By the theory of natural selection, he wrote, all living species have been connected with the parent-species of each genus, by differences not greater than we see between the natural and domestic varieties of the same species at the present day. Thus in the past the number of intermediate and transitional links, between all living and extinct species, must have been inconceivably great. But Darwin knew that the major animal groupswhich modern biologists call phylaappeared fully formed in what were at the time the earliest known fossil-bearing rocks, deposited during a geological period known as the Cambrian. He considered this a serious difficulty for his theory, since if the theory be true, it is indisputable that before the lowest Cambrian stratum was deposited long periods elapsed and that during these vast periods the world swarmed with living creatures. And to the question why we do not find rich fossiliferous deposits belonging to these assumed earliest periods prior to the Cambrian system, I can give no satisfactory answer. So the case at present must remain inexplicable; and may be truly urged as a valid argument against the views here entertained.6

Darwin defended his theory by citing the imperfection of the geological record. In particular, he argued that Precambrian fossils had been destroyed by heat, pressure, and erosion. Some of Darwins modern followers have likewise argued that Precambrian fossils existed but were later destroyed, or that Precambrian organisms were too small or too soft to have fossilized in the first place. Since 1859, however, paleontologists have discovered many Precambrian fossils, many of them microscopic or soft-bodied. As American paleobiologist William Schopf wrote in 1994, The long-held notion that Precambrian organisms must have been too small or too delicate to have been preserved in geological materials [is] now recognized as incorrect. If anything, the abrupt appearance of the major animal phyla about 540 million years agowhich modern biologists call the Cambrian explosion or biologys Big Bangis better documented now than in Darwins time. According to Berkeley paleontologist James Valentine and his colleagues, the explosion is real, it is too big to be masked by flaws in the fossil record. Indeed, as more fossils are discovered it becomes clear that the Cambrian explosion was even more abrupt and extensive than previously envisioned.7

What does Coynes book have to say about this?

Around 600 million years ago, Coyne writes, a whole gamut of relatively simple but multicelled organisms arise, including worms, jellyfish, and sponges. These groups diversify over the next several million years, with terrestrial plants and tetrapods (four-legged animals, the earliest of which were lobe-finned fish) appearing about 400 million years ago.8

In other words, Coynes account of evolutionary history jumps from 600 to 400 million years ago without mentioning the 540 million year-old Cambrian explosion. In this respect, Coynes book reads like a modern biology textbook that has been written to indoctrinate students in Darwinian evolution rather than provide them with the facts.

Coyne goes on to discuss several transitional forms. One of our best examples of an evolutionary transition, he writes, is the fossil record of whales, since we have a chronologically ordered series of fossils, perhaps a lineage of ancestors and descendants, showing their movement from land to water.9

The sequence begins, Coyne writes, with the recently discovered fossil of a close relative of whales, a raccoon-sized animal called Indohyus. Living 48 million years ago, Indohyus was probably very close to what the whale ancestor looked like. In the next paragraph, Coyne writes, Indohyus was not the ancestor of whales, but was almost certainly its cousin. But if we go back 4 million more years, to 52 million years ago, we see what might well be that ancestor. It is a fossil skull from a wolf-sized creature called Pakicetus, which is bit more whalelike than Indohyus. On the page separating these two paragraphs is a figure captioned Transitional forms in the evolution of modern whales, which shows Indohyus as the first in the series and Pakicetus as the second.10

But Pakicetusas Coyne just told usis 4 million years older than Indohyus. To a Darwinist, this doesnt matter: Pakicetus is more whalelike than Indohyus, so it must fall between Indohyus and modern whales, regardless of the fossil evidence.

(Coyne performs the same trick with fossils that are supposedly ancestral to modern birds. The textbook icon Archaeopteryx, with feathered wings like a modern bird but teeth and a tail like a reptile, is dated at 145 million years. But what Coyne calls the nonflying feathered dinosaur fossilswhich should have come before Archaeopteryxare tens of millions of years younger. Like Darwinists Kevin Padian and Luis Chiappe eleven years earlier, Coyne simply rearranges the evidence to fit Darwinian theory.)11

So much for Coynes prediction that later species should have traits that make them look like the descendants of earlier ones. And so much for his argument that if evolution were not true, fossils would not occur in an order that makes evolutionary sense. Ignoring the facts he himself has just presented, Coyne brazenly concludes: When we find transitional forms, they occur in the fossil record precisely where they should. If Coynes book were turned into a movie, this scene might feature Chico Marx saying, Who are you going to believe, me or your own eyes?12

There is another problem with the whale series (and every other series of fossils) that Coyne fails to address: No species in the series could possibly be the ancestor of any other, because all of them possess characteristics they would first have to lose before evolving into a subsequent form. This is why the scientific literature typically shows each species branching off a supposed lineage.

In the figure below, all the lines are hypothetical. The diagram on the left is a representation of evolutionary theory: Species A is ancestral to B, which is ancestral to C, which is ancestral to D, which is ancestral to E. But the diagram on the right is a better representation of the evidence: Species A, B, C and D are not in the actual lineage leading to E, which remains unknown.

It turns out that no series of fossils can provide evidence for Darwinian descent with modification. Even in the case of living species, buried remains cannot generally be used to establish ancestor-descendant relationships. Imagine finding two human skeletons in the same grave, one about thirty years older than the other. Was the older individual the parent of the younger? Without written genealogical records and identifying marks (or in some cases DNA), it is impossible to answer the question. And in this case we would be dealing with two skeletons from the same species that are only a generation apart and from the same location. With fossils from different species that are now extinct, and widely separated in time and space, there is no way to establish that one is the ancestor of anotherno matter how many transitional fossils we find.

In 1978, Gareth Nelson of the American Museum of Natural History wrote: The idea that one can go to the fossil record and expect to empirically recover an ancestor-descendant sequence, be it of species, genera, families, or whatever, has been, and continues to be, a pernicious illusion.13Nature science writer Henry Gee wrote in 1999 that no fossil is buried with its birth certificate. When we call new fossil discoveries missing links, it is as if the chain of ancestry and descent were a real object for our contemplation, and not what it really is: a completely human invention created after the fact, shaped to accord with human prejudices. Gee concluded: To take a line of fossils and claim that they represent a lineage is not a scientific hypothesis that can be tested, but an assertion that carries the same validity as a bedtime storyamusing, perhaps even instructive, but not scientific.14

Embryos

So evolutionary theory needs better evidence than the fossil record can provide. Coyne correctly notes: When he wrote The Origin, Darwin considered embryology his strongest evidence for evolution. Darwin had written that the evidence seemed to show that the embryos of the most distinct species belonging to the same class are closely similar, but become, when fully developed, widely dissimilar, a pattern that reveals community of descent. Indeed, Darwin thought that early embryos show us, more or less completely, the condition of the progenitor of the whole group in its adult state.15

But Darwin was not an embryologist. In The Origin of Species he supported his contention by citing a passage by German embryologist Karl Ernst von Baer:

The embryos of mammals, birds, lizards and snakes, and probably chelonia [turtles] are in their earliest states exceedingly like one another.... In my possession are two little embryos in spirit, whose names I have omitted to attach, and at present I am quite unable to say to what class they belong. They may be lizards or small birds, or very young mammals, so complete is the similarity in the mode of formation of the head and trunk in these animals.16

Coyne claims that this is something von Baer wrote to Darwin, but Coynes history is as unreliable as his paleontology. The passage Darwin cited was from a work written in German by von Baer in 1828; Thomas Henry Huxley translated it into English and published it in 1853. Darwin didnt even realize at first that it was from von Baer: In the first two editions of The Origin of Species he incorrectly attributed the passage to Louis Agassiz.17

Ironically, von Baer was a strong critic of Darwins theory, rejecting the idea that all vertebrates share a common ancestor. According to historian of science Timothy Lenoir, von Baer feared that Darwin and his followers had already accepted the Darwinian evolutionary hypothesis as true before they set to the task of observing embryos. The myth that von Baers work supported Darwins theory was due primarily to another German biologist, Ernst Haeckel.18 Haeckel maintained not only that all vertebrate embryos evolved from a common ancestor, but also that in their development (ontogeny) they replay (recapitulate) their evolutionary history (phylogeny). He called this The Biogenetic Law: Ontogeny recapitulates phylogeny.

In Why Evolution Is True, Coyne writes that the recapitulation of an evolutionary sequence is seen in the developmental sequence of various organs. Each vertebrate undergoes development in a series of stages, and the sequence of those stages happens to follow the evolutionary sequence of its ancestors. The probable reason for this is that as one species evolves into another, the descendant inherits the developmental program of its ancestor. So the descendant tacks changes onto what is already a robust and basic developmental plan. It is best for things that evolved later to be programmed to develop later in the embryo. This adding new stuff onto old principle also explains why the sequence of developmental stages mirrors the evolutionary sequence of organisms. As one group evolves from another, it often adds its developmental program on top of the old one. Thus all vertebrates begin development looking like embryonic fish because we all descended from a fishlike ancestor.19

Nevertheless, Coyne writes, Haeckels Biogenetic Law wasnt strictly true, because embryonic stages dont look like the adult forms of their ancestors, as Haeckel (and Darwin) believed, but like the embryonic forms of their ancestors. But this reformulation of The Biogenetic Law doesnt solve the problem. First, fossil embryos are extremely rare,20 so the reformulated law has to rely on embryos of modern organisms that are assumed to resemble ancestral forms. The result is a circular argument: According to Darwins theory, fish are our ancestors; human embryos (allegedly) look like fish embryos; therefore, human embryos look like the embryos of our ancestors. Theory first, observation laterjust as von Baer had objected.

Second, the idea that later evolutionary stages can simply be tacked onto development is biologically unrealistic. A human is not just a fish embryo with some added features. As British embryologist Walter Garstang pointed out in 1922, a house is not a cottage with an extra story on the top. A house represents a higher grade in the evolution of a residence, but the whole building is alteredfoundations, timbers, and roofeven if the bricks are the same.21

Third, and most important, vertebrate embryos are not most similar in their earliest stages. In the 1860s, Haeckel made some drawings to show that vertebrate embryos look almost identical in their first stagebut his drawings were faked. Not only had he distorted the embryos by making them appear more similar than they really are, but he had also omitted earlier stages in which the embryos are strikingly different from each other. A human embryo in its earliest stages looks nothing like a fish embryo.

Only after vertebrate embryos have progressed halfway through their development do they reach the stage that Darwin and Haeckel treated as the first. Developmental biologists call this different-similar-different pattern the developmental hourglass. Vertebrate embryos do not resemble each other in their earliest stages, but they converge somewhat in appearance midway through development before diverging again. If ontogeny were a recapitulation of phylogeny, such a pattern would be more consistent with separate origins than with common ancestry. Modern Darwinists attempt to salvage their theory by assuming that the common ancestry of vertebrates is obscured because early development can evolve easily, but there is no justification for this assumption other than the theory itself.22

Although Haeckels drawings were exposed as fakes by his own contemporaries, biology textbooks used them throughout the twentieth century to convince students that humans share a common ancestor with fish. Then, in 1997, a scientific journal published an article comparing photos of vertebrate embryos to Haeckels drawings, which the lead author described as one of the most famous fakes in biology. In 2000, Harvard evolutionary biologist Stephen Jay Gould called Haeckels drawings fraudulent and wrote that biologists should be both astonished and ashamed by the century of mindless recycling that has led to the persistence of these drawings in a large number, if not a majority, of modern textbooks.23

But Coyne is not ashamed. He defends Haeckels drawings Haeckel was accused, largely unjustly, Coyne writes, of fudging some drawings of early embryos to make them look more similar than they really are. Yet we shouldnt throw out the baby with the bath water.24 The baby is Darwins theory, which Coyne stubbornly defends regardless of the evidence.

Vestiges and Bad Design

Darwin argued in The Origin of Species that the widespread occurrence of vestigial organsorgans that may have once had a function but are now uselessis evidence against creation. On the view of each organism with all its separate parts having been specially created, how utterly inexplicable is it that organs bearing the plain stamp of inutility should so frequently occur. But such organs, he argued, are readily explained by his theory: On the view of descent with modification, we may conclude that the existence of organs in a rudimentary, imperfect, and useless condition, or quite aborted, far from presenting a strange difficulty, as they assuredly do on the old doctrine of creation, might even have been anticipated in accordance with the views here explained.25

In The Descent of Man, Darwin cited the human appendix as an example of a vestigial organ. But Darwin was mistaken: The appendix is now known to be an important source of antibody-producing blood cells and thus an integral part of the human immune system. It may also serve as a compartment for beneficial bacteria that are needed for normal digestion. So the appendix is not useless at all.26

In 1981, Canadian biologist Steven Scadding argued that although he had no objection to Darwinism, vestigial organs provide no evidence for evolutionary theory. The primarily reason is that it is difficult, if not impossible, to unambiguously identify organs totally lacking in function. Scadding cited the human appendix as an organ previously thought to be vestigial but now known to have a function. Another Canadian biologist, Bruce Naylor, countered that an organ with some function can still be considered vestigial. Furthermore, Naylor argued, perfectly designed organisms necessitated the existence of a creator, but organisms are often something less than perfectly designed and thus better explained by evolution. Scadding replied: The entire argument of Darwin and others regarding vestigial organs hinges on their uselessness and inutility. Otherwise, the argument from vestigiality is nothing more than an argument from homology, and Darwin treated these arguments separately recognizing that they were in fact independent. Scadding also objected that Naylors less than perfectly designed argument was based on a theological assumption about the nature of God, i.e. that he would not create useless structures. Whatever the validity of this theological claim, it certainly cannot be defended as a scientific statement, and thus should be given no place in a scientific discussion of evolution.27

In Why Evolution Is True, Coyne (like Darwin) cites the human appendix as an example of a vestigial organ. Unlike Darwin, however, Coyne concedes that it may be of some small use. The appendix contains patches of tissue that may function as part of the immune system. It has also been suggested that it provides a refuge for useful gut bacteria. But these minor benefits are surely outweighed by the severe problems that come with the human appendix. In any case, Coyne argues, the appendix is still vestigial, for it no longer performs the function for which it evolved.28

As Scadding had pointed out nearly thirty years ago, however, Darwins argument rested on lack of function, not change of function. Furthermore, if vestigiality were redefined as Coyne proposes, it would include many features never before thought to be vestigial. For example, if the human arm evolved from the leg of a four-footed mammal (as Darwinists claim), then the human arm is vestigial. And if (as Coyne argues) the wings of flying birds evolved from feathered forelimbs of dinosaurs that used them for other purposes, then the wings of flying birds are vestigial. This is the opposite of what most people mean by vestigial.29

Coyne also ignores Scaddings other criticism, arguing that whether the human appendix is useless or not, it is an example of imperfect or bad design. What I mean by bad design, Coyne writes, is the notion that if organisms were built from scratch by a designerone who used the biological building blocks or nerves, muscles, bone, and so onthey would not have such imperfections. Perfect design would truly be the sign of a skilled and intelligent designer. Imperfect design is the mark of evolution; in fact, its precisely what we expect from evolution.30

An even better example of bad design, Coyne argues, is the prevalence of dead genes. According to the modern version of Darwinism that Coyne defends, DNA carries a genetic program that encodes proteins that direct embryo development; mutations occasionally alter the genetic program to produce new proteins (or change their locations); and natural selection then sorts those mutations to produce evolution. In the 1970s, however, molecular biologists discovered that most of our DNA does not encode proteins. In 1972 Susumu Ohno called this junk, and in 1976 Richard Dawkins wrote: A large fraction of the DNA is never translated into protein. From the point of view of the individual organism this seems paradoxical. If the purpose of DNA is to supervise the building of bodies, it is surprising to find a large quantity of DNA which does no such thing. From the point of view of Darwinian evolution, however, there is no paradox. The true purpose of DNA is to survive, no more and no less. The simplest way to explain the surplus DNA is to suppose that it is a parasite, or at best a harmless but useless passenger, hitching a ride in the survival machines created by the other DNA.31

Like Dawkins, Coyne regards much of our DNA as parasitic. He writes in Why Evolution Is True: When a trait is no longer used, or becomes reduced, the genes that make it don't instantly disappear from the genome: evolution stops their action by inactivating them, not snipping them out of the DNA. From this we can make a prediction. We expect to find, in the genomes of many species, silenced, or dead, genes: genes that once were useful but are no longer intact or expressed. In other words, there should be vestigial genes. In contrast, the idea that all species were created from scratch predicts that no such genes would exist. Coyne continues: Thirty years ago we couldn't test this prediction because we had no way to read the DNA code. Now, however, its quite easy to sequence the complete genome of species, and its been done for many of them, including humans. This gives us a unique tool to study evolution when we realize that the normal function of a gene is to make a proteina protein whose sequence of amino acids is determined by the sequence of nucleotide bases that make up the DNA. And once we have the DNA sequence of a given gene, we can usually tell if it is expressed normallythat is, whether it makes a functional proteinor whether it is silenced and makes nothing. We can see, for example, whether mutations have changed the gene so that a usable protein can no longer be made, or whether the control regions responsible for turning on a gene have been inactivated. A gene that doesnt function is called a pseudogene. And the evolutionary prediction that well find pseudogenes has been fulfilledamply. Virtually every species harbors dead genes, many of them still active in its relatives. This implies that those genes were also active in a common ancestor, and were killed off in some descendants but not in others. Out of about thirty thousand genes, for example, we humans carry more than two thousand pseudogenes. Our genomeand that of other speciesare truly well populated graveyards of dead genes.32

But Coyne is dead wrong.

Evidence pouring in from genome-sequencing projects shows that virtually all of an organisms DNA is transcribed into RNA, and that even though most of that RNA is not translated into proteins, it performs essential regulatory functions. Every month, science journals publish articles describing more such functions. And this is not late-breaking news: The evidence has been accumulating since 2003 (when scientists finished sequencing the human genome) that pseudogenes and other so-called junk DNA sequences are not useless after all.33Why Evolution Is True ignores this enormous body of evidence, which decisively refutes Coynes Darwinian prediction that our genome should contain lots of dead DNA. Its no wonder that Coyne falls back again and again on the sort of theological arguments that Scadding wrote should be given no place in a scientific discussion of evolution.

Biogeography

Theological arguments are also prominent in The Origin of Species. For example, Darwin argued that the geographic distribution of living things made no sense if species had been separately created, but it did make sense in the context of his theory. Cases such as the presence of peculiar species of bats on oceanic islands and the absence of all other terrestrial mammals, Darwin wrote, are facts utterly inexplicable on the theory of independent acts of creation. In particular: Why, it may be asked, has the supposed creative force produced bats and no other mammals on remote islands? According to Darwin, on my view this question can easily be answered; for no terrestrial mammal can be transported across a wide space of sea, but bats can fly across.34

But Darwin knew that migration cannot account for all patterns of geographic distribution. He wrote in The Origin of Species that the identity of many plants and animals, on mountain-summits, separated from each other by hundreds of miles of lowlands, where Alpine species could not possibly exist, is one of the most striking cases known of the same species living at distant points without the apparent possibility of their having migrated from one point to the other. Darwin argued that the recent ice age affords a simple explanation of these facts. Arctic plants and animals that were nearly the same could have flourished everywhere in Europe and North America, but when the warmth had fully returned, the same species, which had lately lived together on the European and North American lowlands, would again be found in the arctic regions of the Old and New Worlds, and on many isolated mountain-summits far distant from each other.35

So some cases of geographic distribution may not be due to migration, but to the splitting of a formerly large, widespread population into small, isolated populationswhat modern biologists call vicariance. Darwin argued that all modern distributions of species could be explained by these two possibilities. Yet there are many cases of geographic distribution that neither migration nor vicariance seem able to explain.

One example is the worldwide distribution of flightless birds, or ratites. These include ostriches in Africa, rheas in South America, emus and cassowaries in Australia, and kiwis in New Zealand. Since the birds are flightless, explanations based on migration over vast oceanic distances are implausible. After continental drift was discovered in the twentieth century, it was thought that the various populations might have separated with the landmasses. But ostriches and kiwis are much too recent; the continents had already drifted apart when these species originated. So neither migration nor vicariance explain ratite biogeography.36

Another example is freshwater crabs. Studied intensively by Italian biologist Giuseppe Colosi in the 1920s, these animals complete their life cycles exclusively in freshwater habitats and are incapable of surviving prolonged exposure to salt water. Today, very similar species are found in widely separated lakes and rivers in Central and South America, Africa, Madagascar, southern Europe, India, Asia and Australia. Fossil and molecular evidence indicates that these animals originated long after the continents separated, so their distribution is inconsistent with the vicariance hypothesis. Some biologists speculate that the crabs may have migrated by transoceanic rafting in hollow logs, but this seems unlikely given their inability to tolerate salt water. So neither vicariance nor migration provides a convincing explanation for the biogeography of these animals.37

An alternative explanation was suggested in the mid-twentieth century by Lon Croizat, a French biologist raised in Italy. Croizat found that Darwins theory did not seem to agree at all with certain important facts of nature, especially the facts of biogeography. Indeed, he concluded, Darwinism is by now only a straitjacket a thoroughly decrepit skin to hold new wine. Croizat did not argue for independent acts of creation; instead, he proposed that in many cases a widespread primitive species was split into fragments, then its remnants evolved in parallel, in separate locations, into new species that were remarkably similar. Croizat called this process of parallel evolution orthogenesis. Neo-Darwinists such as Ernst Mayr, however, pointed out that there is no mechanism for orthogenesis, which impliescontrary to Darwinismthat evolution is guided in certain directions; so they rejected Croizats hypothesis.38

In Why Evolution Is True, Coyne (like Darwin) attributes the biogeography of oceanic islands to migration, and certain other distributions to vicariance. But Coyne (unlike Darwin) acknowledges that these two processes cannot explain everything. For example, the internal anatomy of marsupial mammals is so different from the internal anatomy of placental mammals that the two groups are thought to have split a long time ago. Yet there are marsupial flying squirrels, anteaters and moles in Australia that strikingly resemble placental flying squirrels, anteaters and moles on other continents, and these forms originated long after the continents had separated.

Coyne attributes the similarities to a well-known process called convergent evolution. According to Coyne. Its really quite simple. Species that live in similar habitats will experience similar selection pressures from their environment, so they may evolve similar adaptations, or converge, coming to look and behave very much alike even though they are unrelated. Put together common ancestry, natural selection, and the origin of species (speciation), add in the fact that distant areas of the world can have similar habitats, and you get convergent evolutionand a simple explanation of a major geographic pattern.39

This is not the same as Croizats orthogenesis, according to which populations of a single species, after becoming separated from each other, evolve in parallel due to some internal directive force. According to Coynes convergent evolution, organisms that are fundamentally different from each other evolve through natural selection to become superficially similar because they inhabit similar environments. The mechanism for orthogenesis is internal, whereas the mechanism for convergence is external. In both cases, however, mechanism is crucial: Without it, orthogenesis and convergence are simply words describing biogeographical patterns, not explanations of how those patterns originated.

So the same question can be asked of convergence that was asked of orthogenesis: What is the evidence for the proposed mechanism? According to Coyne, the mechanism of convergence involves natural selection and speciation.

Selection and Speciation

Coyne writes that Darwin had little direct evidence for selection acting in natural populations. Actually, Darwin had no direct evidence for natural selection; the best he could do in The Origin of Species was give one or two imaginary illustrations. It wasnt until a century later that Bernard Kettlewell provided what he called Darwins missing evidence for natural selectiona shift in the proportion of light- and dark-colored peppered moths that Kettlewell attributed to camouflage and bird predation.40

Since then, biologists have found lots of direct evidence for natural selection. Coyne describes some of it, including an increase in average beak depth of finches on the Galpagos Islands and a change in flowering time in wild mustard plants in Southern Californiaboth due to drought. Like Darwin, Coyne also compares natural selection to the artificial selection used in plant and animal breeding.

But these examples of selectionnatural as well as artificialinvolve only minor changes within existing species. Breeders were familiar with such changes before 1859, which is why Darwin did not write a book titled How Existing Species Change Over Time; he wrote a book titled The Origin of Species by Means of Natural Selection. Darwin called his great work On the Origin of Species, wrote Harvard evolutionary biologist Ernst Mayr in 1982, for he was fully conscious of the fact that the change from one species into another was the most fundamental problem of evolution. Yet, Mayr had written earlier, Darwin failed to solve the problem indicated by the title of his work. In 1997, evolutionary biologist Keith Stewart Thomson wrote: A matter of unfinished business for biologists is the identification of evolution's smoking gun, and the smoking gun of evolution is speciation, not local adaptation and differentiation of populations. Before Darwin, the consensus was that species can vary only within certain limits; indeed, centuries of artificial selection had seemingly demonstrated such limits experimentally. Darwin had to show that the limits could be broken, wrote Thomson, so do we.41

In 2004, Coyne and H. Allen Orr published a detailed book titled Speciation, in which they noted that biologists have not been able to agree on a definition of species because no single definition fits every case. For example, a definition applicable to living, sexually reproducing organisms might make no sense when applied to fossils or bacteria. In fact, there are more than 25 definitions of species. What definition is best? Coyne and Orr argued that, when deciding on a species concept, one should first identify the nature of one's species problem, and then choose the concept best at solving that problem. Like most other Darwinists, Coyne and Orr favor Ernst Mayr's biological species concept (BSC), according to which species are groups of interbreeding natural populations that are reproductively isolated from other such groups. In Why Evolution Is True, Coyne explains that the biological species concept is the one that evolutionists prefer when studying speciation, because it gets you to the heart of the evolutionary question. Under the BSC, if you can explain how reproductive barriers evolve, youve explained the origin of species.42

Theoretically, reproductive barriers arise when geographically separated populations diverge genetically. But Coyne describes five cases of real-time speciation that involve a different mechanism: chromosome doubling, or polyploidy.43 This usually follows hybridization between two existing plant species. Most hybrids are sterile because their mismatched chromosomes cant separate properly to produce fertile pollen and ovaries; occasionally, however, the chromosomes in a hybrid spontaneously double, producing two perfectly matched sets and making reproduction possible. The result is a fertile plant that is reproductively isolated from the two parentsa new species, according to the BSC.

But speciation by polyploidy (secondary speciation) has been observed only in plants. It does not provide evidence for Darwins theory that species originate through natural selection, nor for the neo-Darwinian theory of speciation by geographic separation and genetic divergence. Indeed, according to evolutionary biologist Douglas J. Futuyma, polyploidy does not confer major new morphological characteristics [and] does not cause the evolution of new genera or higher levels in the biological hierarchy.44

So secondary speciation does not solve Darwins problem. Only primary speciationthe splitting of one species into two by natural selectionwould be capable of producing the branching-tree pattern of Darwinian evolution. But no one has ever observed primary speciation. Evolutions smoking gun has never been found.45

Or has it?

In Why Evolution Is True, Coyne claims that primary speciation was observed in an experiment reported in 1998. Curiously, Coyne did not mention it in the 2004 book he co-authored with Orr, but his 2009 account of it is worth quoting in full:

We can even see the origin of a new, ecologically diverse bacterial species, all within a single laboratory flask. Paul Rainey and his colleagues at Oxford University placed a strain of the bacteria Pseudomonas fluorescens in a small vessel containing nutrient broth, and simply watched it. (Its surprising but true that such a vessel actually contains diverse environments. Oxygen concentration, for example, is highest on the top and lowest on the bottom.) Within ten daysno more than a few hundred generationsthe ancestral free-floating smooth bacterium had evolved into two additional forms occupying different parts of the beaker. One, called wrinkly spreader, formed a mat on top of the broth. The other, called fuzzy spreader, formed a carpet on the bottom. The smooth ancestral type persisted in the liquid environment in the middle. Each of the two new forms was genetically different from the ancestor, having evolved through mutation and natural selection to reproduce best in their respective environments. Here, then, is not only evolution but speciation occurring in the lab: the ancestral form produced, and coexisted with, two ecologically different descendants, and in bacteria such forms are considered distinct species. Over a very short time, natural selection in Pseudomonas yielded a small-scale adaptive radiation, the equivalent of how animals or plants form species when they encounter new environments on an oceanic island.46

But Coyne omits the fact that when the ecologically different forms were placed back into the same environment, they suffered a rapid loss of diversity, according to Rainey. In bacteria, an ecologically distinct population (called an ecotype) may constitute a separate species, but only if the distinction is permanent. As evolutionary microbiologist Frederick Cohan wrote in 2002, species in bacteria are ecologically distinct from one another; and they are irreversibly separate.47 The rapid reversal of ecological distinctions when the bacterial populations in Raineys experiment were put back into the same environment refutes Coynes claim that the experiment demonstrated the origin of a new species.

Exaggerating the evidence to prop up Darwinism is not new. In the Galpagos finches, average beak depth reverted to normal after the drought ended. There was no net evolution, much less speciation. Yet Coyne writes in Why Evolution Is True that everything we require of evolution by natural selection was amply documented by the finch studies. Since scientific theories stand or fall on the evidence, Coynes tendency to exaggerate the evidence does not speak well for the theory he is defending. When a 1999 booklet published by The U. S. National Academy of Sciences called the change in finch beaks a particularly compelling example of speciation, Berkeley law professor and Darwin critic Phillip E. Johnson wrote in The Wall Street Journal: When our leading scientists have to resort to the sort of distortion that would land a stock promoter in jail, you know they are in trouble.48

So there are observed instances of secondary speciationwhich is not what Darwinism needsbut no observed instances of primary speciation, not even in bacteria. British bacteriologist Alan H. Linton looked for confirmed reports of primary speciation and concluded in 2001: None exists in the literature claiming that one species has been shown to evolve into another. Bacteria, the simplest form of independent life, are ideal for this kind of study, with generation times of twenty to thirty minutes, and populations achieved after eighteen hours. But throughout 150 years of the science of bacteriology, there is no evidence that one species of bacteria has changed into another.49

Conclusions

Darwin called The Origin of Species one long argument for his theory, but Jerry Coyne has given us one long bluff. Why Evolution Is True tries to defend Darwinian evolution by rearranging the fossil record; by misrepresenting the development of vertebrate embryos; by ignoring evidence for the functionality of allegedly vestigial organs and non-coding DNA, then propping up Darwinism with theological arguments about bad design; by attributing some biogeographical patterns to convergence due to the supposedly well-known processes of natural selection and speciation; and then exaggerating the evidence for selection and speciation to make it seem as though they could accomplish what Darwinism requires of them.

The actual evidence shows that major features of the fossil record are an embarrassment to Darwinian evolution; that early development in vertebrate embryos is more consistent with separate origins than with common ancestry; that non-coding DNA is fully functional, contrary to neo-Darwinian predictions; and that natural selection can accomplish nothing more than artificial selectionwhich is to say, minor changes within existing species.

Faced with such evidence, any other scientific theory would probably have been abandoned long ago. Judged by the normal criteria of empirical science, Darwinism is false. Its persists in spite of the evidence, and the eagerness of Darwin and his followers to defend it with theological arguments about creation and design suggests that its persistence has nothing to do with science at all.50

Nevertheless, biology students might find Coynes book useful. Given accurate information and the freedom to exercise critical thinking, students could learn from Why Evolution Is True how Darwinists manipulate the evidence and mix it with theology to recycle a false theory that should have been discarded long ago.

Notes1 Jerry A. Coyne, Why Evolution Is True (New York: Viking, 2009), p. 3. 2 Coyne, Why Evolution Is True, pp. 3-4. 3 Coyne, Why Evolution Is True, pp. 5-6. 4 Coyne, Why Evolution Is True, pp. 18-19. 5 Coyne, Why Evolution Is True, pp. 17-18, 25. 6 Charles Darwin, The Origin of Species, Sixth Edition (London: John Murray, 1872), Chapter X, pp. 266, 285-288. Available online (2009) here. 7 J. William Schopf, The early evolution of life: solution to Darwins dilemma, Trends in Ecology and Evolution 9 (1994): 375-377. James W. Valentine, Stanley M. Awramik, Philip W. Signor & M. Sadler, The Biological Explosion at the Precambrian-Cambrian Boundary, Evolutionary Biology 25 (1991): 279-356. James W. Valentine & Douglas H. Erwin, Interpreting Great Developmental Experiments: The Fossil Record, pp. 71-107 in Rudolf A. Raff & Elizabeth C. Raff, (editors), Development as an Evolutionary Process (New York: Alan R. Liss, 1987). Jeffrey S. Levinton, The Big Bang of Animal Evolution, Scientific American 267 (November, 1992): 84-91. The Scientific Controversy Over the Cambrian Explosion, Discovery Institute. Available online (2009) here. Jonathan Wells, Icons of Evolution (Washington, DC: Regnery Publishing, 2002), Chapter 3. More information available online (2009) here. Stephen C. Meyer, The Cambrian Explosion: Biologys Big Bang, pp. 323-402 in John Angus Campbell & Stephen C. Meyer (editors), Darwinism, Design, and Public Education (East Lansing, MI: Michigan State University Press, 2003). More information available online (2009) here. 8 Coyne, Why Evolution Is True, p. 28.

9 Coyne, Why Evolution Is True, p. 48. 10 Coyne, Why Evolution Is True, pp. 49-51. 11 Kevin Padian & Luis M. Chiappe, The origin and early evolution of birds, Biological Reviews 73 (1998): 1-42. Available online (2009) here. Wells, Icons of Evolution, pp. 119-122. 12 Coyne, Why Evolution Is True, pp. 25, 53. Chico Marx in Duck Soup (Paramount Pictures, 1933). This and other Marx Brothers quotations are available online (2009) here. 13 Gareth Nelson, Presentation to the American Museum of Natural History (1969), in David M. Williams & Malte C. Ebach, The reform of palaeontology and the rise of biogeography25 years after 'ontogeny, phylogeny, palaeontology and the biogenetic law' (Nelson, 1978), Journal of Biogeography 31 (2004): 685-712. 14 Henry Gee, In Search of Deep Time. New York: Free Press, 1999, pp. 5, 32, 113-117. Jonathan Wells, The Politically Incorrect Guide to Darwinism and Intelligent Design (Washington, DC: Regnery Publishing, 2006). More information available online (2009) here.

15 Coyne, Why Evolution Is True, p. 79. Darwin, The Origin of Species, Chapter XIV, pp. 386-396. Available online (2009) here. 16 Darwin, The Origin of Species, Chapter XIV, pp. 387-388. Available online (2009) here. 17 Coyne, Why Evolution Is True, p. 73. Karl Ernst von Baer, On the Development of Animals, with Observations and Reflections: The Fifth Scholium, translated by Thomas Henry Huxley, pp. 186-237 in Arthur Henfrey & Thomas H. Huxley (editors), Scientific Memoirs: Selected from the Transactions of Foreign Academies of Science and from Foreign Journals: Natural History (London, 1853; reprinted 1966 by Johnson Reprint Corporation, New York). The passage quoted by Darwin is on p. 210. Jane M. Oppenheimer, An Embryological Enigma in the Origin of Species, pp. 221-255 in Jane M. Oppenheimer, Essays in the History of Embryology and Biology (Cambridge, MA: The M.I.T. Press, 1967). 18 Timothy Lenoir, The Strategy of Life (Chicago: The University of Chicago Press, 1982), p. 258. Frederick B. Churchill, The Rise of Classical Descriptive Embryology, pp. 1-29 in Scott F. Gilbert (editor), A Conceptual History of Modern Embryology (Baltimore, MD: The Johns Hopkins University Press, 1991), pp. 19-20. 19 Coyne, Why Evolution Is True, pp. 77-79. 20 Simon Conway Morris, Fossil Embryos, pp. 703-711 in Claudio D. Stern (editor), Gastrulation: From Cells to Embryos (Cold Spring Harbor, NY: Cold Spring Harbor Laboratory Press, 2004). 21 Walter Garstang, The theory of recapitulation: a critical restatement of the biogenetic law, Journal of the Linnean Society (Zoology), 35 (1922): 81-101. 22 See Chapter Five and accompanying references in Wells, Icons of Evolution. See Chapter Three and accompanying references in Wells, The Politically Incorrect Guide to Darwinism and Intelligent Design. 23 Michael K. Richardson, J. Hanken, M. L. Gooneratne, C. Pieau, A. Raynaud, L. Selwood & G. M. Wright, There is no highly conserved embryonic stage in the vertebrates: implications for current theories of evolution and development, Anatomy & Embryology 196 (1997): 91-106. Michael K. Richardson, quoted in Elizabeth Pennisi, Haeckels Embryos: Fraud Rediscovered, Science 277 (1997): 1435. Stephen Jay Gould, Abscheulich! Atrocious! Natural History (March, 2000), pp. 42-49. Hoax of Dodos (2007). Available online (2009) here. 24 Coyne, Why Evolution Is True, p. 78.Notes 25 Darwin, The Origin of Species, Chapters XIV (p. 402) and XV (p. 420). Available online (2009) here. 26 Darwin, Charles. The Descent of Man, First Edition (London: John Murray, 1871), Chapter I (p. 27). Available online (2009) here. Kohtaro Fujihashi, J.R. McGhee, C. Lue, K.W. Beagley, T. Taga, T. Hirano, T. Kishimoto, J. Mestecky & H. Kiyono, Human Appendix B Cells Naturally Express Receptors for and Respond to Interleukin 6 with Selective IgA1 and IgA2 Synthesis, Journal of Clinical Investigations 88 (1991): 248-252. Available online (2009) here. J.A. Laissue, B.B. Chappuis, C. Mller, J.C. Reubi & J.O. Gebbers, The intestinal immune system and its relation to disease, Digestive Diseases (Basel) 11 (1993): 298-312. Abstract available online (2009) here. Loren G. Martin, What is the function of the human appendix? Scientific American (October 21, 1999), Available online (2009) here. R. Randal Bollinger, Andrew S. Barbas, Errol L. Bush, Shu S. Lin & William Parker, Biofilms in the large bowel suggest an apparent function of the human vermiform appendix, Journal of Theoretical Biology 249 (2007): 826-831. Available online (2009) here. Duke University Medical Center, Appendix Isn't Useless At All: It's A Safe House For Good Bacteria, ScienceDaily (October 8, 2007). Available online (2009) here. 27 Steven R. Scadding, Do vestigial organs provide evidence for evolution? Evolutionary Theory 5 (1981): 173-176. Bruce G. Naylor, Vestigial organs are evidence of evolution, Evolutionary Theory 6 (1982): 91-96. Steven R. Scadding, Vestigial organs do not provide scientific evidence for evolution, Evolutionary Theory 6 (1982): 171-173. 28 Coyne, Why Evolution Is True, pp. 61-62. 29 Coyne, Why Evolution Is True, p. 46. 30 Coyne, Why Evolution Is True, pp. 81. 31 Susumu Ohno, So much junk DNA in our genome, Brookhaven Symposia in Biology 23 (1972): 366-70. Richard Dawkins, The Selfish Gene (New York: Oxford University Press, 1976), p. 47. 32 Coyne, Why Evolution Is True, pp. 66-67. 33 A few of the many scientific articles published since 2003 that document the function of so-called junk DNA are: E.S Balakirev & F.J. Ayala, Pseudogenes: are they junk or functional DNA? Annual Review of Genetics 37 (2003): 123-151. A. Httenhofer, P. Schattner & N. Polacek, Non-coding RNAs: hope or hype? Trends in Genetics 21 (2005): 289-297. J.S. Mattick & I.V. Makunin, Non-coding RNA, Human Molecular Genetics 15 (2006): R17-R29. R.K. Slotkin & R. Martienssen, Transposable elements and the epigenetic regulation of the genome, Nature Reviews Genetics 8 (2007): 272-285. P. Carninci, J. Yasuda & Y Hayashizaki, Multifaceted mammalian transcriptome, Current Opinion in Cell Biology 20 (2008): 274-80. C.D. Malone & G.J. Hannon, Small RNAs as Guardians of the Genome, Cell 136 (2009): 656668. C.P. Ponting, P.L. Oliver & W. Reik, Evolution and Functions of Long Noncoding RNAs, Cell 136 (2009): 629641.

34 Darwin, The Origin of Species, Chapters XIII (pp. 347-352) and XV (p. 419). Available online (2009) here. 35 Darwin, The Origin of Species, Chapters XII (pp. 330-332). Available online (2009) here. 36 Alan Cooper, et al., C. Mourer-Chauvir, C.K. Chambers, A. von Haeseler, A.C. Wilson & S. Paabo, Independent origins of New Zealand moas and kiwis, Proceedings of the National Academy of Sciences USA 89 (1992): 8741-8744. Available online (2008) here. Oliver Haddrath & Allan J. Baker, Complete mitochondrial DNA genome sequences of extinct birds: ratite phylogenetics and the vicariance biogeography hypothesis, Proceedings of the Royal Society of London B 268 (2001): 939-945. John Harshman, E.L. Braun, M.J. Braun, C.J. Huddleston, R.C.K. Bowie, J.L. Chojnowski, S.J. Hackett, K.-L. Han, R.T. Kimball, B.D. Marks, K.J. Miglia, W.S. Moore, S. Reddy, F.H. Sheldon, D.W. Steadman, S.J. Steppan, C.C. Witt & T. Yuri, Phylogenomic evidence for multiple losses of flight in ratite birds, Proceedings of the National Academy of Sciences USA 105 (2008): 13462-13467. Abstract available online (2008) here. Giuseppe Sermonti, L'evoluzione in Italia - La via torinese / How Evolution Came to Italy - The Turin Connection, Rivista di Biologia/Biology Forum 94 (2001): 5-12. Available online (2008) here. 37 Giuseppe Colosi, La distribuzione geografica dei Potamonidae, Rivista di Biologia 3 (1921): 294-301. Available online (2009) here. Savel R. Daniels, N. Cumberlidge, M. Prez-Losada, S.A.E. Marijnissen & K.A. Crandall, Evolution of Afrotropical freshwater crab lineages obscured by morphological convergence, Molecular Phylogenetics and Evolution 40 (2006): 227235. Available online (2009) here. R. von Sternberg, N. Cumberlidge & G. Rodriguez. On the marine sister groups of the freshwater crabs (Crustacea: Decapoda: Brachyura), Journal of Zoological Systematics and Evolutionary Research 37 (1999): 1938. Darren C.J. Yeo, et al., Global diversity of crabs (Crustacea: Decapoda: Brachyura) in freshwater, Hydrobiologia 595 (2008): 275-286. 38 Lon Croizat, Space, Time, Form: The Biological Synthesis. Published by the author (Deventer, Netherlands: N. V. Drukkerij Salland, 1962), p. iii. Robin C. Craw, Lon Croizat's Biogeographic Work: A Personal Appreciation, Tuatara 27:1 (August 1984): 8-13. Available online (2009) here. John R. Grehan, Evolution By Law: Croizat's Orthogeny and Darwin's Laws of Growth, Tuatara 27:1 (August 1984): 14-19. Available online (2009) here. Carmen Colacino, Lon Croizats Biogeography and Macroevolution, or Out of Nothing, Nothing Comes, The Philippine Scientist 34 (1997): 73-88. Ernst Mayr, The Growth of Biological Thought (Cambridge, MA: Harvard University Press, 1982), pp. 529-530. 39 Coyne, Why Evolution Is True, pp. 92-94.

40 Coyne, Why Evolution Is True, p. 116. Darwin, The Origin of Species, Chapter IV (p. 70). Available online (2009) here. H. B. D. Kettlewell, Darwins Missing Evidence, Scientific American 200 (March, 1959): 48-53.

41 Ernst Mayr, The Growth of Biological Thought (Cambridge, MA: Harvard University Press, 1982), p. 403. Ernst Mayr, Populations, Species and Evolution (Cambridge, MA: Harvard University Press, 1963), p. 10. Keith Stewart Thomson, Natural Selection and Evolution's Smoking Gun, American Scientist 85 (1997): 516-518.

42 Jerry A. Coyne & H. Allen Orr, Speciation (Sunderland, MA: Sinauer Associates, 2004), p. 25-39. Coyne, Why Evolution Is True, p. 174.

43 Coyne, Why Evolution Is True, p. 188.

44 Douglas J. Futuyma, Evolution (Sunderland, MA: Sinauer Associates, 2005), p. 398.

45 Wells, The Politically Incorrect Guide to Darwinism and Intelligent Design, Chapter Five (The Ultimate Missing Link), pp. 49-59.

46 Coyne, Why Evolution Is True, pp. 129-130.

47 Paul B. Rainey & Michael Travisano. Adaptive radiation in a heterogeneous environment, Nature 394 (1998): 69-72. Frederick M. Cohan, What Are Bacterial Species? Annual Review of Microbiology 56 (2002): 457-482. Available online (2009) here.

48 Coyne, Why Evolution Is True, p. 134. National Academy of Sciences, Science and Creationism: A View from the National Academy of Sciences, Second edition (Washington, DC: National Academy of Sciences Press, 1999), Chapter on Evidence Supporting Biological Evolution, p. 10. Available online (2009) here. Phillip E. Johnson, The Church of Darwin, The Wall Street Journal (August 16, 1999): A14. Available online (2009) here.

49 Alan H. Linton, Scant Search for the Maker, The Times Higher Education Supplement (April 20, 2001), Book Section, p. 29.

Frederick M. Cohan, What Are Bacterial Species? Annual Review of Microbiology 56 (2002): 457-482. Available online (2009) here.

50 Paul A. Nelson, The role of theology in current evolutionary reasoning, Biology and Philosophy 11 (October 1996): 493 - 517. Abstract available online (2009) here. Jonathan Wells, Darwins Straw God Argument, Discovery Institute (December 2008). Available online (2009) here.Jonathan Wells, Darwins Straw God Argument, Discovery Institute (December 2008). Available online (2009) here.

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Modern evolutionary synthesis – Wikipedia

The modern evolutionary synthesis[a] was the widely accepted[1] mid 20th-century synthesis of ideas from fields including genetics, systematics and palaeontology that established evolution as biology's central paradigm.[1][2][3]Embryology was however not integrated into the mid-20th century synthesis; that had to wait for the development of gene manipulation techniques in the 1970s, the growth in understanding of development at a molecular level, and the creation of the modern evolutionary synthesis's successor, evolutionary developmental biology.

The 19th Century ideas of natural selection by Charles Darwin and Mendelian genetics by Gregor Mendel were united by Ronald Fisher, one of the three founders of population genetics, along with J. B. S. Haldane and Sewall Wright, between 1918 and 1932.

The modern synthesis solved difficulties and confusions caused by the specialisation and poor communication between biologists in the early years of the 20th century. At its heart was the question of whether Mendelian genetics could be reconciled with gradual evolution by means of natural selection. A second issue was whether the broad-scale changes of macroevolution seen by palaeontologists could be explained by changes seen in the microevolution of local populations.

The synthesis included evidence from geneticists who studied populations in the field and in the laboratory. These studies were crucial to evolutionary theory. The synthesis drew together ideas from several branches of biology which had become separated, particularly genetics, cytology, systematics, botany, morphology, ecology and paleontology.

Julian Huxley invented the term in his 1942 book, Evolution: The Modern Synthesis. Major figures in the modern synthesis include, Theodosius Dobzhansky, Ivan Schmalhausen,[4]E. B. Ford, Ernst Mayr, Bernhard Rensch, Sergei Chetverikov, George Gaylord Simpson, and G. Ledyard Stebbins.

The modern synthesis of the mid 20th century bridged the gap between the work of experimental geneticists and naturalists, and paleontologists. It states that:[5][6][7]

The idea that speciation occurs after populations are reproductively isolated has been much debated. In plants, polyploidy must be included in any view of speciation. Formulations such as 'evolution consists primarily of changes in the frequencies of alleles between one generation and another' were proposed rather later. The traditional view is that developmental biology played little part in the synthesis,[9] but an account of Gavin de Beer's work by Stephen J. Gould suggests he may be an exception.[10]

Charles Darwin's On the Origin of Species (1859) was successful in convincing most biologists that evolution had occurred, but was less successful in convincing them that natural selection was its primary mechanism. In the 19th and early 20th centuries, variations of Lamarckism, orthogenesis ('progressive' evolution), and saltationism (evolution by jumps) were discussed as alternatives.[11] Also, Darwin did not offer a precise explanation of how new species arise. As part of the disagreement about whether natural selection alone was sufficient to explain speciation, George Romanes coined the term neo-Darwinism to refer to the version of evolution advocated by Alfred Russel Wallace and August Weismann with its heavy dependence on natural selection.[12] Weismann and Wallace rejected the Lamarckian idea of inheritance of acquired characteristics, something that Darwin had not ruled out.[13]

Weismann's idea was that the relationship between the hereditary material, which he called the germ plasm (German, Keimplasma), and the rest of the body (the soma) was a one-way relationship: the germ-plasm formed the body, but the body did not influence the germ-plasm, except indirectly in its participation in a population subject to natural selection. Weismann was translated into English, and though he was influential, it took many years for the full significance of his work to be appreciated.[14] Later, after the completion of the modern synthesis, the term neo-Darwinism came to be associated with its core concept: evolution, driven by natural selection acting on variation produced by genetic mutation, and genetic recombination (chromosomal crossovers).[12]

Gregor Mendel's work was re-discovered by Hugo de Vries and Carl Correns in 1900. News of this reached William Bateson in England, who reported on the paper during a presentation to the Royal Horticultural Society in May 1900.[15] It showed that the contributions of each parent retained their integrity rather than blending with the contribution of the other parent. This reinforced a division of thought, which was already present in the 1890s.[16] The two schools were:

The relevance of Mendelism to evolution was unclear and hotly debated, especially by Bateson, who opposed the biometric ideas of his former teacher Weldon. Many scientists believed the two theories substantially contradicted each other.[18] This debate between the biometricians and the Mendelians continued for some 20 years and was only solved by the development of population genetics.

Thomas Hunt Morgan began his career in genetics as a saltationist, and started out trying to demonstrate that mutations could produce new species in fruit flies. However, the experimental work at his lab with the common fruit fly, Drosophila melanogaster, which helped establish the link between Mendelian genetics and the chromosomal theory of inheritance, demonstrated that rather than creating new species in a single step, mutations increased the genetic variation in the population.[19]

The first step towards the synthesis was the development of population genetics. R. A. Fisher, J. B. S. Haldane, and Sewall Wright provided critical contributions. In 1918, Fisher produced the paper "The Correlation between Relatives on the Supposition of Mendelian Inheritance,"[20] which showed how the continuous variation measured by the biometricians could be the result of the action of many discrete genetic loci. In this and subsequent papers culminating in his 1930 book The Genetical Theory of Natural Selection,[21] Fisher was able to show how Mendelian genetics was, contrary to the thinking of many early geneticists, completely consistent with the idea of evolution driven by natural selection.[22] During the 1920s, a series of papers by Haldane applied mathematical analysis to real-world examples of natural selection such as the evolution of industrial melanism in peppered moths.[22] Haldane established that natural selection could work in the real world at a faster rate than even Fisher had assumed.[23]

Sewall Wright focused on combinations of genes that interacted as complexes, and the effects of inbreeding on small relatively isolated populations, which could exhibit genetic drift. In a 1932 paper, he introduced the concept of an adaptive landscape in which phenomena such as cross breeding and genetic drift in small populations could push them away from adaptive peaks, which would in turn allow natural selection to push them towards new adaptive peaks.[22][24] Wright's model would appeal to field naturalists such as Theodosius Dobzhansky and Ernst Mayr who were becoming aware of the importance of geographical isolation in real world populations.[23] The work of Fisher, Haldane and Wright founded the discipline of population genetics. This is the precursor of the modern synthesis, which is an even broader coalition of ideas.[22][23][25]

Theodosius Dobzhansky, an emigrant from the Soviet Union to the United States, who had been a postdoctoral worker in Morgan's fruit fly lab, was one of the first to apply genetics to natural populations. He worked mostly with Drosophila pseudoobscura. He says pointedly: "Russia has a variety of climates from the Arctic to sub-tropical... Exclusively laboratory workers who neither possess nor wish to have any knowledge of living beings in nature were and are in a minority."[26] Not surprisingly, there were other Russian geneticists with similar ideas, though for some time their work was known to only a few in the West. His 1937 work Genetics and the Origin of Species[27] was a key step in bridging the gap between population geneticists and field naturalists. It presented the conclusions reached by Fisher, Haldane, and especially Wright in their highly mathematical papers in a form that was easily accessible to others. It also emphasized that real world populations had far more genetic variability than the early population geneticists had assumed in their models, and that genetically distinct sub-populations were important. Dobzhansky argued that natural selection worked to maintain genetic diversity as well as driving change. Dobzhansky had been influenced by his exposure in the 1920s to the work of a Russian geneticist Sergei Chetverikov who had looked at the role of recessive genes in maintaining a reservoir of genetic variability in a population before his work was shut down by the rise of Lysenkoism in the Soviet Union.[22][23]

E. B. Ford's work complemented that of Dobzhansky. It was as a result of Ford's work, as well as his own, that Dobzhansky changed the emphasis in the third edition of his famous text from drift to selection.[28] Ford was an experimental naturalist who wanted to test natural selection in nature. He virtually invented the field of research known as ecological genetics. His work on natural selection in wild populations of butterflies and moths was the first to show that predictions made by R. A. Fisher were correct. He was the first to describe and define genetic polymorphism, and to predict that human blood group polymorphisms might be maintained in the population by providing some protection against disease.[29]

Ernst Mayr's key contribution to the synthesis was Systematics and the Origin of Species, published in 1942.[30] Mayr emphasized the importance of allopatric speciation, where geographically isolated sub-populations diverge so far that reproductive isolation occurs. He was skeptical of the reality of sympatric speciation believing that geographical isolation was a prerequisite for building up intrinsic (reproductive) isolating mechanisms. Mayr also introduced the biological species concept that defined a species as a group of interbreeding or potentially interbreeding populations that were reproductively isolated from all other populations.[22][23][31] Before he left Germany for the United States in 1930, Mayr had been influenced by the work of German biologist Bernhard Rensch. In the 1920s Rensch, who like Mayr did field work in Indonesia, analyzed the geographic distribution of polytypic species and complexes of closely related species paying particular attention to how variations between different populations correlated with local environmental factors such as differences in climate. In 1947, Rensch published Neuere Probleme der Abstammungslehre. Die transspezifische Evolution (1959 English translation of 2nd edition: Evolution Above the Species Level).[32] This looked at how the same evolutionary mechanisms involved in speciation might be extended to explain the origins of the differences between the higher level taxa. His writings contributed to the rapid acceptance of the synthesis in Germany.[33][34]

George Gaylord Simpson was responsible for showing that the modern synthesis was compatible with paleontology in his book Tempo and Mode in Evolution published in 1944. Simpson's work was crucial because so many paleontologists had disagreed, in some cases vigorously, with the idea that natural selection was the main mechanism of evolution. It showed that the trends of linear progression (in for example the evolution of the horse) that earlier paleontologists had used as support for neo-Lamarckism and orthogenesis did not hold up under careful examination. Instead the fossil record was consistent with the irregular, branching, and non-directional pattern predicted by the modern synthesis.[22][23]

The botanist G. Ledyard Stebbins extended the synthesis to encompass botany including the important effects of hybridization and polyploidy in plants in his 1950 book Variation and Evolution in Plants.[22]

In 2007, more than half a century after the modern synthesis, Massimo Pigliucci called for an extended evolutionary synthesis to incorporate aspects of biology that had not been included or did not exist in the mid-20th century.[35][36]

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Modern evolutionary synthesis - Wikipedia