Mutationism

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Painting of Hugo de Vries, making a painting of an evening primrose, the plant which had apparently produced new forms by large mutations in his experiments, by Thérèse Schwartze, 1918

 

Mutationism is one of several alternatives to evolution by natural selection that have existed both before and after the publication of Charles Darwin's 1859 book, On the Origin of Species. In the theory, mutation was the source of novelty, creating new forms and new species, potentially instantaneously, in sudden jumps. This was envisaged as driving evolution, which was thought to be limited by the supply of mutations.

Before Darwin, biologists commonly believed in saltationism, the possibility of large evolutionary jumps, including immediate speciation. For example, in 1822 Étienne Geoffroy Saint-Hilaire argued that species could be formed by sudden transformations, or what would later be called macromutation. Darwin opposed saltation, insisting on gradualism in evolution as in geology (uniformitarianism). In 1864, Albert von Kölliker revived Geoffroy's theory. In 1901 the geneticist Hugo de Vries gave the name "mutation" to seemingly new forms that suddenly arose in his experiments on the evening primrose Oenothera lamarckiana, and in the first decade of the 20th century, mutationism, or as de Vries named it mutationstheorie, became a rival to Darwinism supported for a while by geneticists including William Bateson, Thomas Hunt Morgan, and Reginald Punnett.

Understanding of mutationism is clouded by the mid-20th century portrayal of the early mutationists by supporters of the modern synthesis as opponents of Darwinian evolution and rivals of the biometrics school who argued that selection operated on continuous variation. In this portrayal, mutationism was defeated by a synthesis of genetics and natural selection that supposedly started later, around 1918, with work by the mathematician Ronald Fisher. However, the alignment of Mendelian genetics and natural selection began as early as 1902 with a paper by Udny Yule, and built up with theoretical and experimental work in Europe and America. Despite the controversy, the early mutationists had by 1918 already accepted natural selection and explained continuous variation as the result of multiple genes acting on the same characteristic, such as height.

Mutationism, along with other alternatives to Darwinism like Lamarckism and orthogenesis, was discarded by most biologists as they came to see that Mendelian genetics and natural selection could readily work together; mutation took its place as a source of the genetic variation essential for natural selection to work on. However, mutationism did not entirely vanish. In 1940, Richard Goldschmidt again argued for single-step speciation by macromutation, describing the organisms thus produced as "hopeful monsters", earning widespread ridicule. In 1987, Masatoshi Nei argued controversially that evolution was often mutation-limited. Modern biologists such as Douglas J. Futuyma conclude that essentially all claims of evolution driven by large mutations can be explained by Darwinian evolution.

Developments leading up to mutationism

Étienne Geoffroy Saint-Hilaire believed that "monstrosities" could immediately found new species in a single large jump or saltation.

Geoffroy's monstrosities, 1822

Prior to Charles Darwin, most naturalists were saltationists, believing that species evolved and that speciation took place in sudden jumps. Jean-Baptiste Lamarck was a gradualist but similar to other scientists of the period had written that saltational evolution was possible.

In 1822, in the second volume of his Philosophie anatomique, Étienne Geoffroy Saint-Hilaire endorsed a theory of saltational evolution that "monstrosities could become the founding fathers (or mothers) of new species by instantaneous transition from one form to the next." Geoffroy wrote that environmental pressures could produce sudden transformations to establish new species instantaneously.

Darwin's anti-saltationist gradualism, 1859

In his 1859 book On the Origin of Species, Charles Darwin denied saltational evolution. He argued that evolutionary transformation always proceeds gradually, never in jumps: "natural selection acts solely by accumulating slight successive favourable variations, it can produce no great or sudden modification; it can act only by very short steps". Darwin continued in this belief throughout his life.

Rudolph Albert von Kölliker revived Geoffroy's saltationist ideas, calling his theory heterogenesis. It depended on a nonmaterial directive force (orthogenesis).

 

Thomas Henry Huxley warned Darwin that he had taken on "an unnecessary difficulty in adopting Natura non facit saltum ["Nature does not take leaps"] so unreservedly." Huxley feared this assumption could discourage naturalists (catastrophists) who believed that major leaps and cataclysms played a significant role in the history of life.

von Kölliker's heterogenesis, 1864

In 1864 Albert von Kölliker revived Geoffroy's theory that evolution proceeds by large steps, under the name of heterogenesis, but this time assuming the influence of a nonmaterial force to direct the course of evolution.

Galton's "sports", 1892

Darwin's cousin, Francis Galton, considered Darwin's evidence for evolution, and came to an opposite conclusion about the type of variation on which natural selection must act. He carried out his own experiments and published a series of papers and books setting out his views. Already by 1869 when he published Hereditary Genius, he believed in evolution by saltation. In his 1889 book Natural Inheritance he argued that natural selection would benefit from accepting that the steps need not, as Darwin had stated, be minute. In his 1892 book Finger Prints, he stated directly that "The progress of evolution is not a smooth and uniform progression, but one that proceeds by jerks, through successive 'sports' (as they are called), some of them implying considerable organic changes; and each in its turn being favoured by Natural Selection".

From 1860 to 1880 saltation had been a minority viewpoint, to the extent that Galton felt his writings were being universally ignored. By 1890 it became a widely held theory, and his views helped to launch a major controversy.

Drawing of William Bateson, 1909, by the biologist Dennis G. Lillie

Bateson's discontinuous variation, 1894

William Bateson's 1894 book Materials for the Study of Variation, Treated with Especial Regard to Discontinuity in the Origin of Species marked the arrival of mutationist thinking, before the rediscovery of Mendel's laws. He examined discontinuous variation (implying a form of saltation) where it occurred naturally, following William Keith Brooks, Galton, Thomas Henry Huxley and St. George Jackson Mivart.

Early 20th century mutationism

De Vries and Mendelian mutationstheorie, 1901

The main principle of the mutation theory is that species and varieties have originated by mutation, but are, at present, not known to have originated in any other way. — Hugo de Vries

Hugo de Vries's careful 1901 studies of wild variants of the evening primrose Oenothera lamarckiana showed that distinct new forms could arise suddenly in nature, apparently at random, and could be propagated for many generations without dissipation or blending. He gave such changes the name "mutation". By this, de Vries meant that a new form of the plant was created in a single step (not the same as a mutation in the modern sense); no long period of natural selection was required for speciation, and nor was reproductive isolation. In the view of the historian of science Peter J. Bowler, De Vries used the term to mean

large-scale genetic changes capable of producing a new subspecies, or even species, instantaneously.

The historian of science Betty Smocovitis described mutationism as

the case of purported saltatory evolution that Hugo de Vries had mistakenly interpreted for the evening primrose, Oenothera.

De Vries set out his position, known as Mutationstheorie (mutation theory) on the creative nature of mutation in his 1905 book Species and Varieties: their Origin by Mutation. In the view of the historian of science Edward Larson, de Vries was the person largely responsible for transforming Victorian era saltationism into early 20th century mutation theory, "and in doing so pushed Darwinism near the verge of extinction as a viable scientific theory".

Johannsen's "pure line" experiments, 1903

Wilhelm Johannsen's "pure line" experiments seemed to show that evolution could not work on continuous variation.

 

In the early 1900s, Darwin's mechanism of natural selection was understood by believers in continuous variation, principally the biometricians Walter Weldon and Karl Pearson, to be able to work on a continuously varying characteristic, whereas de Vries argued that selection on such characteristics would be ineffective. Wilhelm Johannsen's "pure line" experiments on Phaseolus vulgaris beans appeared to refute this mechanism. Using the true-breeding Princess variety of bean, carefully inbred within weight classes, Johannsen's work appeared to support de Vries. The offspring had a smooth random distribution. Johanssen believed that his results showed that continuous variability was not inherited, so evolution must rely on discontinuous mutations, as de Vries had argued. Johanssen published his work in Danish in a 1903 paper Om arvelighed i samfund og i rene linier (On inheritance in populations and in pure lines), and in his 1905 book Arvelighedslærens Elementer (The Elements of Heredity).

Punnett's mimicry, 1915

Papilio polytes has 3 forms with differing wing patterns, here the "Romulus" morph. Reginald Punnett argued that this polymorphism demonstrated discontinuous evolution. However, Ronald Fisher showed that this could have arisen by small changes in additional modifier genes.

 

In 1915, Reginald Punnett argued in his book Mimicry in Butterflies that the 3 morphs (forms) of the butterfly Papilio polytes, which mimic different host species of butterfly, demonstrated discontinuous evolution in action. The different forms existed in a stable polymorphism controlled by 2 Mendelian factors (genes). The alleles of these genes were certainly discontinuous, so Punnett supposed that they must have evolved in discontinuous leaps.

The undermining of mutationism

Yule's analysis of Mendelism and continuous variation, 1902

The undermining of mutationism began almost at once, in 1902, as the statistician Udny Yule analysed Mendel's theory and showed that given full dominance of one allele over another, a 3:1 ratio of alleles would be sustained indefinitely. This meant that the recessive allele could remain in the population with no need to invoke mutation. He also showed that given multiple factors, Mendel's theory enabled continuous variation, as indeed Mendel had suggested, removing the central plank of the mutationist theory, and criticised Bateson's confrontational approach. However, the "excellent" paper did not prevent the Mendelians and the biometricians from falling out.

Nilsson-Ehle's experiments on Mendelian inheritance and continuous variation, 1908

The Swedish geneticist H. Nilsson-Ehle demonstrated in 1908, in a paper published in German in a Swedish journal, Einige Ergebnisse von Kreuzungen bei Hafer und Weizen (Observations on Crosses in Oats and Wheat), that continuous variation could readily be produced by multiple Mendelian genes. He found numerous Mendelian 3:1 ratios, implying a dominant and a recessive allele, in oats and wheat; a 15:1 ratio for a cross of oat varieties with black and white glumes respectively, implying two pairs of alleles (two Mendelian factors); and that crossing a red-grained Swedish velvet wheat with a white one gave in the third (F3) generation the complex signature of ratios expected of three factors at once, with 37 grains giving only red offspring, 8 giving 63:1 in their offspring, 12 giving 15:1, and 6 giving 3:1. There weren't any grains giving all white, but as he had only expected 1 of those in his sample, 0 was not an unlikely outcome. Genes could clearly combine in almost infinite combinations: ten of his factors allowed for almost 60,000 different forms, with no need to suppose that any new mutations were involved. The results implied that natural selection would work on Mendelian genes, helping to bring about the unification of Darwinian evolution and genetics.

Similar work in America by Edward East on maize in 1910 showed the same thing for biologists without access to Nilsson-Ehle's work. On the same theme, the mathematician Ronald Fisher published "The Correlation Between Relatives on the Supposition of Mendelian Inheritance" in 1918, again showing that continuous variation could readily be produced by multiple Mendelian genes. It showed, too, that Mendelian inheritance had no essential link with mutationism: Fisher stressed that small variations (per gene) would be sufficient for natural selection to drive evolution.

Castle's selection experiments on hooded rats, 1911

Starting in 1906, William Castle carried out a long study of the effect of selection on coat colour in rats. The piebald or hooded pattern was recessive to the grey wild type. He crossed hooded rats with the black-backed Irish type, and then back-crossed the offspring with pure hooded rats. The dark stripe on the back was bigger. He then tried selecting different groups for bigger or smaller stripes for 5 generations, and found that it was possible to change the characteristics way beyond the initial range of variation. This effectively refuted de Vries's claim that continuous variation could not be inherited permanently, requiring new mutations. By 1911 Castle noted that the results could be explained by Darwinian selection on heritable variation of Mendelian genes.

Morgan's small Mendelian genes in Drosophila, 1912

Thomas Hunt Morgan's work on Drosophila melanogaster found many small Mendelian factors for natural selection to work on.

 

By 1912, after years of work on the genetics of Drosophila fruit flies, Thomas Hunt Morgan showed that these animals had many small Mendelian factors on which Darwinian evolution could work as if variation was fully continuous. The way was open for geneticists to conclude that Mendelism supported Darwinism.

Muller's balanced lethal explanation of Oenothera "mutations", 1918

De Vries's mutationism was dealt a serious if not fatal blow in 1918 by the American geneticist Hermann Joseph Muller. He compared the behaviour of balanced lethals in Drosophila with De Vries's supposed mutations in Oenothera, showing that they could work the same way. No actual mutations were involved, but infrequent chromosome crossovers accounted for the sudden appearance of traits which had been present in the genes all along.

Fisher's explanation of polymorphism, 1927

In 1927, Fisher explicitly attacked Punnett's 1915 theory of discontinuous evolution of mimicry. Fisher argued that selection acting on genes making small modifications to the butterfly's phenotype (its appearance) would allow the multiple forms of a polymorphism to be established.

Later mutationist theories

The understanding that Mendelian genetics could both preserve discrete variations indefinitely, and support continuous variation for natural selection to work on gradually, meant that most biologists from around 1918 onwards accepted natural selection as the driving force of evolution. Mutationism and other alternatives to evolution by natural selection did not however vanish entirely.

Berg's nomogenesis, 1922

Lev Berg proposed a combination of mutationism and directed (orthogenetic) evolution in his 1922 book Nomogenesis; or, Evolution Determined by Law. He used evidence from paleontology, zoology, and botany to argue that natural selection had limitations which set a direction for evolution. He claimed that speciation was caused by "mass transformation of a great number of individuals" by directed mass mutations.

John Christopher Willis's The Course of Evolution by Differentiation Or Divergent Mutation Rather Than by Selection, 1940

Willis's macromutations, 1923

In 1923, the botanist John Christopher Willis proposed that species were formed by large mutations, not gradual evolution by natural selection, and that evolution was driven by orthogenesis, which he called "differentiation", rather than by natural selection.

Goldschmidt's hopeful monsters, 1940

Masatoshi Nei argues that evolution is often mutation-limited.

 

In his 1940 book The Material Basis of Evolution, the German geneticist Richard Goldschmidt argued for single-step speciation by macromutation, describing the organisms thus produced as "hopeful monsters". Goldschmidt's thesis was universally rejected and widely ridiculed by biologists, who favoured the neo-Darwinian explanations of Fisher, J. B. S. Haldane and Sewall Wright. However, interest in Goldschmidt's ideas has reawakened in the field of evolutionary developmental biology.

Nei's mutation-driven evolution, 1987

Contemporary biologists accept that mutation and selection both play roles in evolution; the mainstream view is that while mutation supplies material for selection in the form of variation, all non-random outcomes are caused by natural selection. Masatoshi Nei argues instead that the production of more efficient genotypes by mutation is fundamental for evolution, and that evolution is often mutation-limited. Nei's book received thoughtful reviews; while Wright, in the conservative journal Evolution, rejected Nei's thinking as mistaken, Galtier, Weiss, Stoltzfus, and Wagner, although not necessarily agreeing with Nei's position, treated it as a relevant alternative view.

Contemporary approaches

Reviewing the history of macroevolutionary theories, the American evolutionary biologist Douglas J. Futuyma notes that since 1970, two very different alternatives to Darwinian gradualism have been proposed, both by Stephen Jay Gould: mutationism, and punctuated equilibria. Gould's macromutation theory gave a nod to his predecessor with an envisaged "Goldschmidt break" between evolution within a species and speciation. His advocacy of Goldschmidt was attacked with "highly unflattering comments" by Brian Charlesworth, and Alan Templeton. Futuyma concludes, following other biologists reviewing the field such as K.Sterelny and A. Minelli, that essentially all the claims of evolution driven by large mutations could be explained within the Darwinian evolutionary synthesis. James A. Shapiro's claim that molecular genetics undermines Darwinism has been described as mutationism and an extreme view by the zoologist Andy Gardner.

Multiple explanations have been offered since the 19th century for how evolution took place, given that many scientists initially had objections to natural selection. Many of these, including mutationism, led to some form of orthogenesis (solid blue arrows), with or without theistic intervention (dotted blue arrows). Better understanding of mutation led instead (dashed orange arrows) to the modern synthesis of Mendelian genetics with natural selection, establishing Darwinian evolution throughout biology.

Historiography

Biologists at the start of the 20th century broadly agreed that evolution occurred, but felt that the mechanisms suggested by Darwin, including natural selection, would be ineffective. Large mutations looked likely to drive evolution quickly, and avoided the difficulty which had rightly worried Darwin, namely that blending inheritance would average out any small favourable changes. Further, large saltatory mutation, able to create species in a single step, offered a ready explanation of why the fossil record should contain large discontinuities and times of rapid change.

These discoveries were often framed by supporters of the mid-20th century modern synthesis, such as Julian Huxley and Ernst Mayr, as a controversy between the early geneticists—the "Mendelians"—including Bateson, Johannsen, de Vries, Morgan, and Punnett, who advocated Mendelism and mutation, and were understood as opponents of Darwin's original gradualist view, and the biometricians such as Pearson and Weldon, who opposed Mendelism and were more faithful to Darwin. In this version, little progress was made during the eclipse of Darwinism, and the debate between mutationist geneticists such as de Vries and biometricians such as Pearson ended with the victory of the modern synthesis between about 1918 and 1950. According to this account, the new population genetics of the 1940s demonstrated the explanatory power of natural selection, while mutationism, alongside other non-Darwinian approaches such as orthogenesis and structuralism, was essentially abandoned. This view became dominant in the second half of the 20th century, and was accepted by both biologists and historians.

A more recent view, advocated by the historians Arlin Stoltzfus and Kele Cable, is that Bateson, de Vries, Morgan and Punnett had by 1918 formed a synthesis of Mendelism and mutationism. The understanding achieved by these geneticists spanned the action of natural selection on alleles (alternative forms of a gene), the Hardy-Weinberg equilibrium, the evolution of continuously-varying traits (like height), and the probability that a new mutation will become fixed. In this view, the early geneticists accepted natural selection alongside mutation, but rejected Darwin's non-Mendelian ideas about variation and heredity, and the synthesis began soon after 1900. The traditional claim that Mendelians rejected the idea of continuous variation outright is simply false; as early as 1902, Bateson and Edith Saunders wrote that "If there were even so few as, say, four or five pairs of possible allelomorphs, the various homo- and hetero-zygous combinations might, on seriation, give so near an approach to a continuous curve, that the purity of the elements would be unsuspected".

Historians have interpreted the history of mutationism in different ways.The classical view is that mutationism, opposed to Darwin's gradualism, was an obvious error; the decades-long delay in synthesizing genetics and Darwinism is an "inexplicable embarrassment"; genetics led logically to the modern synthesis and mutationism was one of several anti-Darwinian "blind alleys" separate from the main line leading from Darwin to the present. A revisionist view is that mutationists accepted both mutation and selection, with broadly the same roles they have today, and early on accepted and indeed offered a correct explanation for continuous variation based on multiple genes, paving the way for gradual evolution. At the time of the Darwin centennial in Cambridge in 1909, mutationism and Lamarckism were contrasted with natural selection as competing ideas; 50 years later, at the 1959 University of Chicago centennial of the publication of On the Origin of Species, mutationism was no longer seriously considered.

History of zoology since 1859

From Wikipedia, the free encyclopedia

This article considers the history of zoology since the theory of evolution by natural selection proposed by Charles Darwin in 1859.

Charles Darwin gave new direction to morphology and physiology, by uniting them in a common biological theory: the theory of organic evolution. The result was a reconstruction of the classification of animals upon a genealogical basis, fresh investigation of the development of animals, and early attempts to determine their genetic relationships. The end of the 19th century saw the fall of spontaneous generation and the rise of the germ theory of disease, though the mechanism of inheritance remained a mystery. In the early 20th century, the rediscovery of Mendel's work led to the rapid development of genetics by Thomas Hunt Morgan and his students, and by the 1930s the combination of population genetics and natural selection in the "neo-Darwinian synthesis".

Second half of nineteenth century

Darwin and the theory of evolution

The 1859 publication of Darwin's theory in On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life is often considered the central event in the history of modern zoology. Darwin's established credibility as a naturalist, the sober tone of the work, and most of all the sheer strength and volume of evidence presented, allowed Origin to succeed where previous evolutionary works such as the anonymous Vestiges of Creation had failed. Most scientists were convinced of evolution and common descent by the end of the 19th century. However, natural selection would not be accepted as the primary mechanism of evolution until well into the 20th century, as most contemporary theories of heredity seemed incompatible with the inheritance of random variation.

Alfred Russel Wallace, following on earlier work by de Candolle, Humboldt and Darwin, made major contributions to zoogeography. Because of his interest in the transmutation hypothesis, he paid particular attention to the geographical distribution of closely allied species during his field work first in South America and then in the Malay archipelago. While in the archipelago he identified the Wallace line, which runs through the Spice Islands dividing the fauna of the archipelago between an Asian zone and a New Guinea/Australian zone. His key question, as to why the fauna of islands with such similar climates should be so different, could only be answered by considering their origin. In 1876 he wrote The Geographical Distribution of Animals, which was the standard reference work for over half a century, and a sequel, Island Life, in 1880 that focused on island biogeography. He extended the six-zone system developed by Philip Sclater for describing the geographical distribution of birds to animals of all kinds. His method of tabulating data on animal groups in geographic zones highlighted the discontinuities; and his appreciation of evolution allowed him to propose rational explanations, which had not been done before.

The scientific study of heredity grew rapidly in the wake of Darwin's Origin of Species with the work of Francis Galton and the biometricians. The origin of genetics is usually traced to the 1866 work of the monk Gregor Mendel, who would later be credited with the laws of inheritance. However, his work was not recognized as significant until 35 years afterward. In the meantime, a variety of theories of inheritance (based on pangenesis, orthogenesis, or other mechanisms) were debated and investigated vigorously.

In 1859, Charles Darwin placed the whole theory of organic evolution on a new footing, by his discovery of a process by which organic evolution can occur, and provided observational evidence that it had done so. This changed the attitudes of most exponents of the scientific method. Darwin's discoveries revolutionised the zoological and botanical sciences, by introducing the theory of evolution by natural selection as an explanation for the diversity of all animal and plant life. The subject-matter of this new science, or branch of biological science, had been neglected: it did not form part of the studies of the collector and systematist, nor was it a branch of anatomy, nor of the physiology pursued by medical men, nor again was it included in the field of microscopy and the cell theory. Almost a thousand years before Darwin, the Arab scholar Al-Jahiz (781 – 868) had already developed a rudimentary theory of natural selection , describing the Struggle for existence in his Book of Animals where he speculates on how environmental factors can affect the characteristics of species by forcing them to adapt and then passing on those new traits to future generations. However, his work had largely been forgotten, along with many other early advances of Arab scientists, and there is no evidence that his works were known to Darwin. 

The area of biological knowledge which Darwin was the first to subject to scientific method and to render, as it were, contributory to the great stream formed by the union of the various branches, is that which relates to the breeding of animals and plants, their congenital variations, and the transmission and perpetuation of those variations. This branch of biological science may be called thremmatology - the science of breeding. Outside the scientific world, an immense mass of observation and experiment had grown up in relation to this subject. From the earliest times the shepherd, the farmer, the horticulturist, and the fancier had for practical purposes made themselves acquainted with a number of biological laws, and successfully applied them without exciting more than an occasional notice from the academic students of biology. Darwin made use of these observations and formulated their results to a large extent as the laws of variation and heredity. As the breeder selects a congenital variation which suits his requirements, and by breeding from the animals (or plants) exhibiting that variation obtains a new breed specially characterised by that variation, so in nature is there a selection amongst all the congenital variations of each generation of a species. This selection depends on the fact that more young are born than the natural provision of food will support. In consequence of this excess of births there is a struggle for existence and a survival of the fittest, and consequently an ever-present necessarily acting selection, which either maintains accurately the form of the species from generation to generation or leads to its modification in correspondence with changes in the surrounding circumstances which have relation to its fitness for success in the struggle for life, structures to the service of the organisms in which they occur.

It cannot be said that previously to Darwin there had been any very profound study of teleology, but it had been the delight of a certain type of mind, that of the lovers of nature or naturalists par excellence as they were sometimes termed, to watch the habits of living animals and plants and to point out the remarkable ways in which the structure of each variety of organic life was adapted to the special circumstances of life of the variety or species. The astonishing colours and grotesque forms of some animals and plants which the museum zoologists gravely described without comment were shown by these observers of living nature to have their significance in the economy of the organism possessing them; and a general doctrine was recognized, to the effect that no part or structure of an organism is without definite use and adaptation, being designed by the Creator for the benefit of the creature to which it belongs, or else for the benefit, amusement or instruction of his highest creatureman. Teleology in this form of the doctrine of design was never very deeply rooted amongst scientific anatomists and systematists. It was considered permissible to speculate somewhat vaguely on the subject of the utility of this or that startling variety of structure; but few attempts, though some of great importance, were made systematically to explain by observation and experiment the adaptation of organic structures to particular purposes in the case of the lower animals and plants. Teleology had, indeed, an important part in the development of physiology - the knowledge of the mechanism, the physical and chemical properties, of the parts of the body of man and the higher animals allied to him. But, as applied to lower and more obscure forms of life, teleology presented almost insurmountable difficulties; and consequently, in place of exact experiment and demonstration, the most reckless though ingenious assumptions were made as to the utility of the parts and organs of lower animals.

Darwin's theory had as one of its results the reformation and rehabilitation of teleology. According to that theory, every organ, every part, colour and peculiarity of an organism, must either be of benefit to that organism itself or have been so to its ancestors: no peculiarity of structure or general conformation, no habit or instinct in any organism, can be supposed to exist for the benefit or amusement of another organism, not even for the delectation of man himself.

A very subtle and important qualification of this generalization has to be recognized (and was recognized by Darwin) in the fact that owing to the interdependence of the parts of the bodies of living things and their profound chemical interactions and peculiar structural balance (what is called organic polarity) the variation of one single part (a spot of colour, a tooth, a claw, a leaflet) may, and demonstrably does in many cases entail variation of other parts what are called correlated variations. Hence many structures which are obvious to the eye, and serve as distinguishing marks of separate species, are really not themselves of value or use, but are the necessary concomitants of less obvious and even altogether obscure qualities, which are the real characters upon which selection is acting. Such correlated variations may attain to great size and complexity without being of use. But eventually they may in turn become, in changed conditions, of selective value. Thus in many cases the difficulty of supposing that selection has acted on minute and imperceptible initial variations, so small as to have no selective value, may be got rid of. A useless correlated variation may have attained great volume and quality before it is (as it were) seized upon and perfected by natural selection. All organisms are essentially and necessarily built up by such correlated variations.

Necessarily, according to the theory of natural selection, structures either are present because they are selected as useful or because they are still inherited from ancestors to whom they were useful, though no longer useful to the existing representatives of those ancestors. Structures previously inexplicable were now explained as survivals from a past age, no longer useful though once of value. Every variety of form and colour was urgently and absolutely called upon to produce its title to existence either as an active useful agent or as a survival. Darwin himself spent a large part of the later years of his life in thus extending the new teleology.

The old doctrine of types, which was used by the philosophically minded zoologists (and botanists) of the first half of the 19th century as a ready means of explaining the failures and difficulties of the doctrine of design, fell into its proper place under the new dispensation. The adherence to type, the favourite conception. of the transcendental morphologist, was seen to be nothing more than the expression of one of the laws of thremmatology, the persistence of hereditary transmission of ancestral characters, even when they have ceased to be significant or valuable in the struggle for existence, whilst the so-called evidences of design which was supposed to modify the limitations of types assigned to Himself by the Creator were seen to be adaptations due to the selection and intensification by selective breeding of fortuitous congenital variations, which happened to prove more useful than the many thousand other variations which did not survive in the struggle for existence.

Thus not only did Darwins theory give a new basis to the study of organic structure, but, whilst rendering the general theory of organic evolution equally acceptable and necessary, it explained the existence of low and simple forms of life as survivals of the earliest ancestry of the more highly complex forms, and revealed the classifications of the systematist as unconscious attempts to construct the genealogical tree or pedigree of plants and animals. Finally, it brought the simplest living matter or formless protoplasm before the mental vision as the starting point whence, by the operation of necessary mechanical causes, the highest forms have been evolved, and it rendered unavoidable the conclusion that this earliest living material was itself evolved by gradual processes, the result also of the known and recognized laws of physics and chemistry, from material which we should call not living. It abolished the conception of life as an entity above and beyond the common properties of matter, and led to the conviction that the marvellous and exceptional qualities of that which we call living matter are nothing more nor less than an exceptionally complicated development of those chemical and physical properties which we recognize in a gradually ascending scale of evolution in the carbon compounds, containing nitrogen as well as oxygen, sulphur and hydrogen as constituent atoms of their enormous molecules. Thus mysticism was finally banished from the domain of biology, and zoology became one of the physical sciencesthe science which seeks to arrange and discuss the phenomena of animal life and form, as the outcome of the operation of the laws of physics and chemistry.

A subdivision of zoology which was at one time in favour is simply into morphology and physiology, the study of form and structure on the one hand, and the study of the activities and functions of the forms and structures of the other. But a logical division like this is not necessarily conducive to the ascertainment and remembrance of the historical progress and present significance of the science. No such distinction of mental activities as that involved in the division of the study of animal life into morphology and physiology has ever really existed: the investigator of animal forms has never entirely ignored the functions of the forms studied by him, and the experimental inquirer into the functions and properties of animal tissues and organs has always taken very careful account of the forms of those tissues and organs. A more instructive subdivision must be one which corresponds to the separate currents of thought and mental preoccupation which have been historically manifested in western Europe in the gradual evolution of what is to-day the great river of zoological doctrine to which they have all been rendered contributory.

Cell theory, embryology and germ theory

Innovative experimental methods such as Louis Pasteur's contributed to the young field of bacteriology in the late 19th century.

 

Cell theory led zoologists to re-envision individual organisms as interdependent assemblages of individual cells. Scientists in the rising field of cytology, armed with increasingly powerful microscopes and new staining methods, soon found that even single cells were far more complex than the homogeneous fluid-filled chambers described by earlier microscopists. Much of the research on cell reproduction came together in August Weismann's theory of heredity: he identified the nucleus (in particular chromosomes) as the hereditary material, proposed the distinction between somatic cells and germ cells (arguing that chromosome number must be halved for germ cells, a precursor to the concept of meiosis), and adopted Hugo de Vries's theory of pangenes. Weismannism was extremely influential, especially in the new field of experimental embryology.

By the 1880s, bacteriology was becoming a coherent discipline, especially through the work of Robert Koch, who introduced methods for growing pure cultures on agar gels containing specific nutrients in Petri dishes. The long-held idea that living organisms could easily originate from nonliving matter (spontaneous generation) was attacked in a series of experiments carried out by Louis Pasteur, while debates over vitalism vs. mechanism (a perennial issue since the time of Aristotle and the Greek atomists) continued apace.

Physiology

Over the course of the 19th century, the scope of physiology expanded greatly, from a primarily medically oriented field to a wide-ranging investigation of the physical and chemical processes of life—including plants, animals, and even microorganisms in addition to man. Living things as machines became a dominant metaphor in biological (and social) thinking. Physiologists such as Claude Bernard explored (through vivisection and other experimental methods) the chemical and physical functions of living bodies to an unprecedented degree, laying the groundwork for endocrinology (a field that developed quickly after the discovery of the first hormone, secretin, in 1902), biomechanics, and the study of nutrition and digestion. The importance and diversity of experimental physiology methods, within both medicine and zoology, grew dramatically over the second half of the 19th century. The control and manipulation of life processes became a central concern, and experiment was placed at the center of biological education.

Twentieth century

At the beginning of the 20th century, zoological research was largely a professional endeavour. Most work was still done in the natural history mode, which emphasized morphological and phylogenetic analysis over experiment-based causal explanations. However, anti-vitalist experimental physiologists and embryologists, especially in Europe, were increasingly influential. The tremendous success of experimental approaches to development, heredity, and metabolism in the 1900s and 1910s demonstrated the power of experimentation in biology. In the following decades, experimental work replaced natural history as the dominant mode of research.

Early 20th century work (variation and heredity)

After publication of his work The Origin of Species, Darwin became interested in the animal and plant mechanisms that confer advantages to individual members of a species. Much important work was done by Fritz Muller (Für Darwin), by Hermann Müller (Fertilization of Plants by Insects), August Weismann, Edward B. Poulton and Abbott Thayer. There was considerable progress during this period in the field that would become known as genetics, the laws of variation and heredity (originally known as thremmatology). The progress of microscopy gave a clearer understanding of the origin of the egg-cell and sperm-cell and the process of fertilization.

Mendel and zoology

Mendel's experiments on cultivated varieties of plants were published in 1865, but attracted little notice until thirty-five years later, sixteen years after his death. Mendel tried to gain a better understanding of heredity. His main experiments were with varieties of the edible pea. He chose a variety with one marked structural feature and crossed it with another variety in which that feature was absent. For example, he hybridized a tall variety, with a dwarf variety, a yellow-seeded variety with a green-seeded variety, and a smooth-seeded variety with a wrinkle-seeded variety. In each experiment, he concentrated on one character; after obtaining a first hybrid generation, he allowed the hybrids to self-fertilize, and recorded the number of individuals in the first, second, third, and fourth generations in which the chosen character appeared.

In the first hybrid generation, nearly all the individuals had the positive character, but in subsequent generations the positive character was not present in all individuals: half had the character and half did not. Thus the random pairing of two groups of reproductive cells yielded the proportion 1 PP, 2 PN, 1 NN, where P stands for the character and N for its absence - the character was present in three-quarters of the offspring and absent from a quarter. The failure of the character to distribute itself among all of the reproductive cells of a hybrid individual, and the limitation of its distribution to half only of those cells, prevents the swamping of a new character by interbreeding. The tendency of the proportions in the offspring is to give, in a series of generations, a reversion from the hybrid form PN to a race with the positive character and a race without it. This tendency favours the persistence of a new character of large volume suddenly appearing in a stock. The observations of Mendel thus favoured the view that the variations upon which natural selection acts are not small but large and discontinuous. However, it did not appear that large variations would be favoured any more than small ones, or that the eliminating action of natural selection upon an unfavourable variation could be checked.

Much confusion arose in discussions of this topic, because of defective nomenclature. Some authors used the word mutation only for large variations that appeared suddenly and that could be inherited, and fluctuation for small variations, whether they could be transmitted or not. Other authors used fluctuation only for small, acquired variations due to changes in food, moisture and other features of the environment. This kind of variation is not heritable, but the small variations Darwin thought important are. The best classification of the variations in organisms separates those that arise from congenital variations from those that arise from variations of the environment or the food-supply. The former are innate variations, the latter are "acquired variations". Both innate and acquired variations include some that are more and some that are less obvious. There are slight innate variations in every new generation of every species; their greatness or smallness so far as human perception goes is not of much significance, their importance for the origin of new species depends on whether they are valuable to the organism in the struggle for existence and reproduction. An imperceptible physiological difference might be of selective value, and it might carry with it correlated variations that may or may not appeal to the human eye, but are of no selective value themselves.

The views of Hugo de Vries and others about the importance of saltatory variation, the soundness of which was still not generally accepted in 1910, may be gathered from the article Mendelism. A due appreciation of the far-reaching results of correlated variation must, it appeared, give a new and distinct explanation of large mutations, discontinuous variation, and saltatory evolution. The analysis of the specific variations of organic form to determine the nature and limitation of a single character, and whether two variations of a structural unit can blend when one is transmitted by the male parent and the other by the female, were yet to be determined. It was not clear whether absolute blending was possible, or whether all apparent blending was only a more-or-less minutely subdivided mosaic of non-combinable characters of the parents.

Another important development of Darwin's conclusions deserves notice. The fact of variation was familiar: no two animals, even of the same brood, are alike. Jean-Baptiste Lamarck hypothesised that structural alterations acquired by a parent might be transmitted to the offspring, and as these are acquired by an animal or plant as a consequence of the action of the environment, the offspring would sometimes start with a greater fitness for those conditions than its parents started with. In turn, it would acquire a greater development of the same modification, which it would transmit to its offspring. Lamarck argued that, over several generations, a structural alteration might thus be acquired. The familiar illustration of Lamarck's hypothesis is that of the giraffe, whose long neck might, he suggested, was acquired by the efforts of a short-necked race of herbivores who stretched their necks to reach the foliage of trees in a land where grass was deficient, the effort producing a longer neck of each generation, which was then transmitted to the next. This process is known as 'direct adaptation'.

Such structural adaptations are acquired by an animal in the course of its life, but are limited in degree and rare, rather than frequent and obvious. Whether acquired characters could be transmitted to the next generation was a very different issue. Darwin excluded any assumption of the transmission of acquired characters. He pointed to the fact of congenital variation, and showed that congenital variations are arbitrary and non-significant.

Congenital variation

At the beginning of the 20th century, the causes of congenital variation were obscure, although it was recognised that they were largely due to a mixing of the matter that constituted the fertilized germ or embryo-cell from two individuals. Darwin had shown that congenital variation was all-important. A popular illustration of the difference was this: a man born with four fingers only on his right hand might transmit this peculiarity to at least some of his children; but a man with one finger chopped off will produce children with five fingers. Darwin, influenced by some facts that seemed to favour the Lamarckian hypothesis, thought that acquired characters are sometimes transmitted, but did not consider that this mechanism was likely to be of great importance.

After Darwin's writings, there was an effort to find evidence for the transmission of acquired characters; ultimately, the Lamarckian hypothesis of transmission of acquired characters was not supported by evidence, and was dismissed. August Weismann argued from the structure of the egg-cell and sperm-cell, and from how and when they are derived in the growth of the embryo from the egg, that it was impossible that a change in parental structure could produce a representative change in the germ or sperm-cells.

The only evidence that seemed to support the Lamarckian hypothesis were the experiments of Charles Brown-Séquard, who produced epilepsy in guinea-pigs by bisection of the large nerves or spinal cord, which led him to believe that, in rare instances, the artificially-produced epilepsy and mutilation of the nerves was transmitted. The record of Brown-Séquard's original experiments was unsatisfactory, and attempts reproduce them were unsuccessful. Conversely, the vast number of experiments in the cropping of the tails and ears of domestic animals, as well as of similar operations on man, had negative results. Stories of tailess kittens, puppies, and calves, born from parents one of whom had been thus injured, are abundant, but failed to stand experimental examination.

Whilst evidence of the transmission of an acquired character proved wanting, the a priori arguments in its favour were recognized as flawed, and cases that appeared to favour the Lamarckian assumption were found to be better explained by the Darwinian principle. For example, the occurrence of blind animals in caves and in the deep sea was a fact that even Darwin regarded as best explained by the atrophy of the eye in successive generations through the absence of light and consequent disuse. However, it was suggested that this is better explained by natural selection acting on congenital fortuitous variations. Some animals are born with distorted or defective eyes. If a number of some species of fish are swept into a cavern, those with perfect eyes would follow the light and eventually escape, leaving behind those with imperfect eyes to breed in the dark place. In every succeeding generation this would be the case, and even those with weak but still seeing eyes would escape, until only a pure race of blind animals would be left in the cavern.

Transmission

It was argued that the elaborate structural adaptations of the nervous system that underlie instincts must have been slowly built up by the transmission to offspring of acquired experience. It seemed hard to understand how complicated instincts could be due to the selection of congenital variations, or be explained except by the transmission of habits acquired by the parent. However, imitation of the parent by the young account for some, and there are cases in which elaborate actions must be due to the natural selection of a fortuitously-developed habit. Such cases are the habits of 'shamming dead' and the combined posturing and colour peculiarities of certain caterpillars (Lepidoptera larvae) that cause them to resemble dead twigs or similar objects. The advantage to the caterpillar is that it escapes (say) a bird that would, were it not deceived, attack and eat it. Preceding generations of caterpillars cannot have acquired this habit of posturing by experience; either a caterpillar postures and escapes, or it does not posture and is eaten - it is not half eaten and allowed to profit by experience. Thus, we seem justified in assuming that there are many movements of stretching and posturing possible to caterpillars, that some had a fortuitous tendency to one position, some to another, and, that among all the variety of habitual movements, one is selected and perpetuated because it happened to make the caterpillar look more like a twig.

Record of the past

Man, compared with other animals, has the fewest instincts and the largest brain in proportion to body size. He builds up, from birth onwards, his own mental mechanisms, and forms more of them, and takes longer in doing so, than any other animal. The later stages of evolution from ape-like ancestors have consisted in the acquisition of a larger brain and in the education of that brain. A new feature in organic development makes its appearance when we set out the facts of man's evolutionary history. This factor is the record of the past, which grows and develops by laws other than those affecting the perishable bodies of successive generations of mankind, so that man, by the interaction of the record and his educability, is subject to laws of development unlike those by which the rest of the living world is governed.

Ecology and environmental science

In the early 20th century, naturalists were faced with increasing pressure to add rigor and preferably experimentation to their methods, as the newly prominent laboratory-based biological disciplines had done. Ecology had emerged as a combination of biogeography with the biogeochemical cycle concept pioneered by chemists; field biologists developed quantitative methods such as the quadrat and adapted laboratory instruments and cameras for the field to further set their work apart from traditional natural history. Zoologists did what they could to mitigate the unpredictability of the living world, performing laboratory experiments and studying semi-controlled natural environments; new institutions like the Carnegie Station for Experimental Evolution and the Marine Biological Laboratory provided more controlled environments for studying organisms through their entire life cycles.

Charles Elton's studies of animal food chains was pioneering among the succession of quantitative methods that colonized the developing ecological specialties. Ecology became an independent discipline in the 1940s and 1950s after Eugene P. Odum synthesized many of the concepts of ecosystem ecology, placing relationships between groups of organisms (especially material and energy relationships) at the center of the field. In the 1960s, as evolutionary theorists explored the possibility of multiple units of selection, ecologists turned to evolutionary approaches. In population ecology, debate over group selection was brief but vigorous; by 1970, most zoologists agreed that natural selection was rarely effective above the level of individual organisms.

Classical genetics, the modern synthesis, and evolutionary theory

Thomas Hunt Morgan's illustration of crossing over, part of the Mendelian-chromosome theory of heredity

 

1900 marked the so-called rediscovery of Mendel: Hugo de Vries, Carl Correns, and Erich von Tschermak independently arrived at Mendel's laws (which were not actually present in Mendel's work). Soon after, cytologists (cell biologists) proposed that chromosomes were the hereditary material. Between 1910 and 1915, Thomas Hunt Morgan and the "Drosophilists" in his fly lab forged these two ideas—both controversial—into the "Mendelian-chromosome theory" of heredity. They quantified the phenomenon of genetic linkage and postulated that genes reside on chromosomes like beads on string; they hypothesized crossing over to explain linkage and constructed genetic maps of the fruit fly Drosophila melanogaster, which became a widely used model organism.

Hugo de Vries tried to link the new genetics with evolution; building on his work with heredity and hybridization, he proposed a theory of mutationism, which was widely accepted in the early 20th century. Lamarckism also had many adherents. Darwinism was seen as incompatible with the continuously variable traits studied by biometricians, which seemed only partially heritable. In the 1920s and 1930s—following the acceptance of the Mendelian-chromosome theory— the emergence of the discipline of population genetics, with the work of R.A. Fisher, J.B.S. Haldane and Sewall Wright, unified the idea of evolution by natural selection with Mendelian genetics, producing the modern synthesis. The inheritance of acquired characters was rejected, while mutationism gave way as genetic theories matured.

In the second half of the century the ideas of population genetics began to be applied in the new discipline of the genetics of behavior, sociobiology, and, especially in humans, evolutionary psychology. In the 1960s W.D. Hamilton and others developed game theory approaches to explain altruism from an evolutionary perspective through kin selection. The possible origin of higher organisms through endosymbiosis, and contrasting approaches to molecular evolution in the gene-centered view (which held selection as the predominant cause of evolution) and the neutral theory (which made genetic drift a key factor) spawned perennial debates over the proper balance of adaptationism and contingency in evolutionary theory.

In the 1970s Stephen Jay Gould and Niles Eldredge proposed the theory of punctuated equilibrium which holds that stasis is the most prominent feature of the fossil record, and that most evolutionary changes occur rapidly over relatively short periods of time. In 1980 Luis Alvarez and Walter Alvarez proposed the hypothesis that an impact event was responsible for the Cretaceous–Paleogene extinction event. Also in the early 1980s, statistical analysis of the fossil record of marine organisms published by Jack Sepkoski and David M. Raup lead to a better appreciation of the importance of mass extinction events to the history of life on earth.

Twenty-first century

Advances were made in analytical chemistry and physics instrumentation including improved sensors, optics, tracers, instrumentation, signal processing, networks, robots, satellites, and compute power for data collection, storage, analysis, modeling, visualization, and simulations. These technology advances allowed theoretical and experimental research including internet publication of zoological science. This enabled worldwide access to better measurements, theoretical models, complex simulations, theory predictive model experimentation, analysis, worldwide internet observational data reporting, open peer-review, collaboration, and internet publication.