Darwin, Charles Robert
Charles Robert Darwin. Wikimedia Commons (Public Domain)

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Darwin, Charles Robert

Darwin, Charles Robert

(b. The Mount, Shrewsbury, England, 9 February 1809; d. Down House, Downe, Kent, England, 19 April 1882)

natural history, geology, evolution.

Darwin was the fifth child and second son of Robert Waring Darwin, a successful and respected physician practicing in Shrewsbury, and Susannah Wedgwood, daughter of the potter Josiah Wedgwood I. The Darwin family was characterized by the high intellectual quality of its members, prosperity, industriousness, professional ability, and wide cultural interests.

Darwin’s education was begun by his elder sisters after the premature death of his mother. In 1817 he was sent to a day school at Shrewsbury, where he was found to be slow at learning. In 1818 he was entered at Shrewsbury School under Dr. Samuel Butler (grandfather of the author of Erewhon). He afterwards complained he was taught nothing but classics, a little ancient history, and geography. “The school as a means of education to me was simply a blank.” This statement is perhaps unfair in that his grounding in the classics probably helped him afterwards to think so devastatingly straight. But his headmaster rebuked him publicly for wasting his time on chemical experiments, and his father upbraided him with the remark, “You care for nothing but shooting, dogs, and ratcatching, and you will be a disgrace to yourself and all your family.” He was removed from Shrewsbury School in 1825 and sent to Edinburgh University to study medicine.

Darwin stayed at Edinburgh until 1827, attending courses of lectures on materia medica, pharmacy, chemistry, and anatomy, which he found unbearably dull. But worst of all were his experiences attending operations, performed perforce without anesthetics. These repelled him to such an extent that he rushed out and vowed that medicine was no career for him. The chief advantages that he gained from his stay at Edinburgh were his friendship with Robert Grant, zoologist, who accepted Lamarck’s teachings on evolution; geological excursions with Robert Jameson; and expeditions onto the Firth of Forth to collect marine animals.

Confronted with the necessity of starting his son on a new line of endeavor, Darwin’s father sent him to Cambridge as a preparation for entering the Church of England as a clergyman. At that time, he accepted the Articles of Faith, and perfunctorily attended lectures, but his three years at Cambridge he thought “wasted as far as the academical studies were concerned, as completely as at Edinburgh and at school.” But he there made the acquaintance of Adam Sedgwick, who interested him in geology, and, most important of all, John Stevens Henslow, who fired him with a passionate interest in natural history and inspired him with a friendship that gave him confidence in himself, after his discouragement in his own family, at school, and at Edinburgh. Henslow acted as a second father to young Darwin during this critical, formative period in his life, which was climaxed by the voyage of the Beagle.

After taking a poor degree at Cambridge in 1831, Darwin was at home when he received an invitation, instigated by Henslow, to join the Admiralty survey ship H. M. S. Beagle, under the command of Robert FitzRoy, as unpaid naturalist on a voyage to survey the coasts of Patagonia, Tierra del Fuego, Chile, and Peru; to visit some Pacific islands; and to carry a chain of chronometrical stations around the world. Darwin wished to accept at once, but his father objected. Darwin’s uncle, Josiah Wedgwood II, thought the objection unreasonable and persuaded his brother-in-law to withdraw it. Darwin sailed on the Beagle on 27 December 1831.

The five years of the voyage were the most important event in Darwin’s intellectual life and in the history of biological science. Darwin sailed with no formal scientific training. He returned a hard-headed man of science, knowing the importance of evidence, almost convinced that species had not always been as they were since the creation but had undergone change. He also developed doubts of the value of the Scriptures as a trustworthy guide to the history of the earth and of man, with the result that he gradually became an agnostic. The experiences of his five years in the Beagle, how he dealt with them, and what they led to, built up into a process of epoch-making importance in the history of thought.

On 29 January 1839, Darwin married his first cousin, Emma Wedgwood, daughter of the Josiah Wedgwood who had saved his place in the Beagle. At first the young couple lived in London, but ill health began to attack Darwin shortly after his marriage. He found himself increasingly handicapped by great lassitude, nausea, and intestinal discomfort, which made it impossible for him to lead a normal life with social and academic contacts. The Darwins therefore moved to the country, fifteen miles from London, to Down House, in the village of Downe, in Kent. There Darwin had a garden, a small estate about which he could walk, spare rooms for the few friends who were occasionally invited to stay, a study where he could work, and peace and quiet.

His ill health gradually restricted him to a routine of four hours’ work a day, walks in the garden, visits to the greenhouses, occasional walks in the neighborhood or rides on a pony, interspersed with periods of rest on a sofa while reading a novel. Dinner was followed by listening to the piano and then early bed.

As his physicians could not discover any organic cause for Darwin’s ill health, the idea gained ground that he was a hypochondriac, and in recent years some psychiatrists have attempted to show that he was of a neurotic disposition, but the evidence is not wholly satisfactory. Following the suggestion of Saul Adler, a pathologist expert on South American infections, other commentators note that Darwin was heavily bitten in the Argentine by the Benchuca, the “black bug of the pampas,” Triatoma infestans, 70 percent of which are now reckoned to be vectors of Trypanosoma cruzi, the causative agent of Chagas’ disease. As this trypanosome was not discovered until 1909, Darwin’s doctors would have been unable to find an organic cause for his trouble; but the clinical picture found in sufferers from Chagas’ disease matches Darwin’s symptoms in detail. The parasite invades the muscle of the heart causing lassitude; it also invades the nerve cells of Auerbach’s plexus in the wall of the intestine, upsetting peristalsis; and it invades the auricular-ventricular bundle of the heart, interference with which may cause heart block. It has unpredictable periods of latency, after which it can be recovered from blood of patients infected many years before. Darwin suffered heart attacks and died from one. Whether in addition Darwin showed neurotic tendencies, accepting the overcare which his wife bestowed on him and taking advantage of his semi-invalidism to protect himself from the distractions of society and amusement, will never be known.

Darwin and his wife had ten children, of whom three died in infancy or childhood. The healths of the survivors (all of whom lived to ripe ages), and their launching in life, gave him anxiety. He once wrote that his chief worries were the discovery of gold in California and in Australia, “beggaring me by making my money on mortgage worthless”; the French coming by the Westerham and Sevenoaks roads, and, therefore, enclosing Down; and, third, “professions for my boys.” His fears were groundless. His investments were very profitable, Napoleon III did not invade England, and his sons made careers for themselves of which any father could be proud: Sir George Darwin, mathematician and astronomer; Sir Francis Darwin, botanist; Leonard Darwin, engineer and promoter of eugenics; and Sir Horace Darwin, scientific engineer and founder of the Cambridge Scientific Instrument Co. These men, their children and grandchildren, like their father and ancestors, provide material for Galton’s theory of the inheritability of intellectual eminence.

Darwin’s contributions to science fall into three main groups: geology, evolution and natural selection, and botany.

Darwin as Geologist. When Darwin sailed in the Beagle, the most commonly accepted theory in geology was that of catastrophism, which held that the forces which had produced change during the past history of the earth had been on a far greater scale than any which are active today. When geologists had concluded that stratified rocks had originated as beds of sediment laid down beneath the sea, they had then been obliged to explain how the sedimentary strata had been raised up from the sea bottom to form hills and mountains, and how they had been shifted from the horizontal position, in which they must originally have been deposited, to the inclined or even vertical positions in which they are now so often found. There were two possible explanations: (1) that the sea had formerly stood at a higher level and had in fact extended over the whole earth’s surface, or (2) that the strata had been upraised and disturbed. By 1831 most geologists had rejected the idea that the sea had formerly covered the whole earth, but when they tried to imagine forces capable of raising rock strata from the bottom of the sea to form hills or ranges of mountains, it seemed to them that such changes would necessarily involve tremendous convulsions of the earth’s surface. Furthermore, they saw a link between periods of convulsion and upheaval in the history of the earth and changes in animal life. They concluded, naturally, that a convulsion of the earth’s surface tended to destroy many species of animals and that when the convulsion had subsided new species were created to replace those that had been lost.

In the view of the catastrophists, there had been in the history of the earth a succession of creations of animal and plant life. Each creation had survived during a period of relative tranquillity of the earth’s surface, but had later been destroyed in a catastrophic disturbance and upheaval. The history of the earth therefore included periods of tranquillity interrupted by catastrophes, and each catastrophe was followed by a new creation. The catastrophic theory of geological history was reconciled to the account of creation in the Bible by supposing that the most recent catastrophe had been the flood of Noah. Many fossil animals were therefore characterized as antediluvian, or from before the flood. Geologists also supposed that the successive creations were of a progressively higher order which culminated finally in the creation of man.

Since the time of the flood the surface of the earth had been fixed, stable, and unchanging and the whole system of nature tranquil and orderly. The modern order of nature, catastrophists thought, was so very different from that which had existed throughout most of the history of the earth that little could be inferred of past events from present conditions.

The catastrophist viewpoint, firmly held by such English geologists as William Buckland, William Daniel Conybeare, and Adam Sedgwick, was challenged in 1830 by Charles Lyell in his Principles of Geology. Lyell had concluded that strata could be elevated from the bottom of the sea by the repeated action of earthquakes continued over long periods of time. In Italy and Sicily he had found recently elevated strata in the neighborhood of active volcanoes and in an area where severe earthquakes had occurred frequently throughout historic time. He showed to his own satisfaction that the strata had been elevated very gradually over an immense period of time, instead of by a single convulsion, and then went on to investigate how far the phenomena of geology might be explained by the various forces observable on the earth’s surface today. With great success he showed that the ordinary action of rain, running water, and the waves of the sea was sufficient to explain the wearing down of land and deposition of sediments, whereas the ordinary, long-continued action of volcanoes and earthquakes was sufficient to elevate continents and mountain ranges.

When Darwin sailed in the Beagle, he had with him the first volume of Lyell’s Principles of Geology (1830), which he had been advised to read but on no account to believe. That is, however, exactly what Darwin did, because he satisfied himself that Lyell’s views accorded with the facts: the first step in the making of a man of science possessed of critical judgment.

Santiago in the Cape Verde Islands provided Darwin’s first material for geological study. By applying Lyell’s principles, he quickly unraveled the history of the island. From the clear exposure of the rocks on the coast he saw that the oldest were crystalline and volcanic. Overlying them was a bed of limestone containing shells of marine organisms of Tertiary date, sixty feet above sea level and twenty feet thick. This bed, Darwin reasoned, had been deposited in a shallow sea and had since been raised to its present height; the island had undergone subsidence and then elevation. Above the limestone was more recent volcanic rock which had altered the uppermost limestone and baked it; evidence of metamorphism in situ.

St. Paul’s Rocks in mid-Atlantic enabled him to make acquaintance with an island which he correctly recognized as being of neither volcanic nor coralline origin; it is regarded today as an exposed portion of the mantle of the earth. But it was South America that presented itself as a model for his observational and interpretative skill. By comparing lavas of undoubted volcanic origin with igneous rocks of the Andes, he saw that they were closely related. He found that minerals in granites and in lavas were in all respects similar, and he showed a perfect graded series between crystalline granites and glasslike lavas. By observing the direction of strike and the angle of dip of strata, he found that the planes of foliation in schists and gneisses and the planes of cleavage in slates remained constant over areas stretching hundreds of miles, while their planes of inclination could vary greatly. Foliation planes and cleavage planes were parallel to the direction of the great axes along which elevation of the land had taken place. This was directly contrary to the then accepted views put forward by Sedgwick and by Lyell, and Darwin was able to prove that foliation and cleavage are not original phenomena dependent on the deposition of the strata, but had been subsequently superimposed on the beds by pressure, resulting in recrystallization in the case of foliation. He observed clay slate passing over into gneiss as it approached granite, which showed how homogeneous rocks could develop foliation by metamorphosis. Darwin’s demonstration of the origin of metamorphic rocks by deformation and of the distinction between cleavage and sedimentary bedding was a major contribution to geology.

The west coast of South America was an equally fruitful geological laboratory for Darwin. He was a witness to the earthquake that devastated Concepción and saw that it had resulted in an elevation of the land by several feet. He was further able to make out a connection between elevation of the land and neighboring volcanic activity because at the time when the land rose near Concepción a number of volcanoes in the Andes renewed their activity, and a new volcano beneath the sea erupted near Juan Fernandez. All these facts were shortly to be woven into an unexpected pattern.

By observing that the shells in beds raised to heights of as much as 1,300 feet belonged to species still living, and in proportions of species similar to those in the ocean, Darwin was able to prove, as Lyell had in Sicily, that such elevation was recent. In the Andes, at a height of 7,000 feet, he found a fossil forest of trees in situ (i. e., not drifted from elsewhere) because their trunks projected several feet perpendicularly from the stratum in which they were embedded, which was itself overlaid by thousands of feet of sedimentary deposits alternating with volcanic lavas. This showed that the trees had been buried thousands of feet beneath sea level as the deposits accumulated, and had since been raised to their present height. Here was proof of extensive subsidence as well as elevation.

Beds containing shells that had been raised up were far fewer and much less extensive than low-lying beds, and Darwin found that this was due to the erosion that constantly attacks land surfaces and destroys deposits. This fact had a bearing on estimations of the age of the earth, because while the time taken for a deposit to be formed can be roughly gauged, it is impossible to tell how long an interval of time has gone by while the beds were eroded. This means that the nonconformities between geological formations may represent much longer periods of time than the formation of the beds themselves, and that the age of the earth must be vastly greater than was then imagined.

From these findings, Darwin was led to consider what are the most favorable conditions for the accumulation of beds many thousands of feet thick containing fossils, such as are found elsewhere. He concluded that it could only have been when a slow subsidence continually lowered the bottom of a shallow sea, so that the depth of that sea below sea level remained fairly constant. These conditions further imply that such a sea was near to continental land, from which erosion brought down the material spread out in the deposits. Proof of this was obtained in observation of beds south of Valparaiso, 800 feet thick, containing shells of animals which when alive lived in shallow water, not more than 100 feet deep. The lowest beds of the deposit must have lain 700 feet higher than at present when the first shells became embedded, and as subsidence started the deposits continued and kept pace with the drop of 700 feet. Another bed showed evidence of the same process involving a subsidence and deposits of 6,000 feet. The various conditions required for such accumulations of fossils can only have been met occasionally and at particular localities in the history of the earth, from which it follows that the fossil record must be expected to be erratic and incomplete.

Another line of study to which Darwin was led by his expeditions in the Andes was the change in climate which regions must have undergone, both in geological time and during the life of man. This followed from observation of deserted houses, abandoned by Indians, up to the snowline in places now quite uninhabitable because of lack of water and fertility. This connection between geological phenomena and life was further developed in his studies of coral reefs.

While in South American waters, Darwin had never seen a coral reef, but he knew that atolls in their myriads all over the oceans are about at sea level, and that the coral polyps which build them can live only in waters less than 120 feet deep and at temperatures above 20°C. The theory of the origin of atolls, or lagoon islands, developed by Lyell, was that they were formed on the rims of submarine volcanic craters. Darwin quickly saw that such a hypothesis was untenable, and he put forward his own deductively. There must have been a foundation for the corals at a depth not exceeding 120 feet. It cannot be imagined that there were innumerable submarine mountain ranges with their summits all exactly at this height; therefore, they must have been brought to the required level by subsidence, and he found evidence for it in drowned villages in the Caroline Islands. This has since been confirmed by deep borings at Funafuti, at Bikini, and at Eniwetok, where evidence has been found of over 4,600 feet of coral rock produced by shallow-water organisms during prolonged subsidence.

In addition to atolls or lagoon-islands, there are also barrier reefs, like the Great Barrier Reef of Australia, which is known to have been erected by coral polyps on the down-faulted coastal edge of northeast Australia. Darwin observed a third type of coral reef, the fringing reefs or shore-reefs that surround an island which projects high above the sea, and these reefs are often found lifted above sea level. Fringing reefs are therefore the result of coral action on platforms that are either stationary or have undergone elevation. With all this evidence, Darwin drew a map. He found that large zones of the Pacific Ocean have atolls and barrier reefs and have undergone subsidence, while other zones, parallel to these, have fringing reefs and have undergone elevation. Now the most extraordinary correlation with these facts, bearing in mind Darwin’s observations of the connection between land elevation and volcanic activity, was the distribution of active volcanoes in the Pacific: all of them are situated in the zone of fringing reefs, not one of them in the zone of atolls or barrier reefs. Darwin’s book on Coral Reefs, published in 1842, still remains the accepted explanation, except for a slight addition in the form of R. A. Daly’s glacial control theory. This is based on the fact that during the recent Ice Ages, the amount of water immobilized in icecaps reduced the level of the oceans by 150 feet, which would have exposed the atolls to the air and cut down their height by wave-action. But when the Ice Age came to an end and the sea level rose again, the platforms were there for coral polyps to make new atolls.

The evidence which Darwin obtained in South America and in atolls for changes of sea level was later responsible for leading him into error. In 1838 in attempting to explain the Parallel Roads of Glen Roy in Scotland, Darwin imagined that the land had sunk by over 1,000 feet and that the so-called roads were marine beaches. In 1862 it was shown by T. F. Jamieson that the glaciers of Ben Nevis had dammed back the waters of a lake, the level of which dropped twice as outlets were formed, and that the roads were lake beaches. Darwin admitted defeat in words which have an important message for all scientific research workers: “How rash it is in science to argue because any case is not one thing, it must be some second thing, which happens to be known to the writer…. My error has been a good lesson to me never to trust in science to the principle of exclusion”.

Darwin’s trilogy of geological books was completed by the publication in 1844 of Volcanic Island, and at the end of 1846 by Geological Observations on South America. The supremacy of the position which geology occupied in Darwin’s mind shortly after his return from the voyage of the Beagle is shown by the order of subjects in the title of the first edition, published in August 1839, of his Journal of Researches into the Geology and Natural History of the Various Countries Visited by H. M. S. Beagle. In the second edition, published in August 1845, natural history and geology have changed places. The reason for this is to be found in the progressive orientation of Darwin’s mind toward the problem of evolution.

Evolution and Natural Selection. When Darwin sailed in the Beagle, he had no reason to call in question the accepted view that the species of plants and animals alive on earth were as they had always been since the creation. It was only gradually that doubts began. They arose from four kinds of evidence. The first was that in some areas species had become extinct. Darwin himself had found gigantic fossil armadillos (and other forms) in South America; but armadillos of similar but not identical form also live in South America. This meant that “extinct animals have a close relation in form with extinct species.” Why was this?

Second, in adjacent areas of South America, Darwin found one species replaced by different, although very similar, species. On the pampas he observed and collected specimens of the South American ostrich (rhea), but when he went farther south into Patagonia, he found a very similar but smaller species (Rhea darwinii). But why were the two species of rheas built on the same plan, different from that of the African ostriches? Why were the agoutis and vizcachas of South America built on the same plan as other South American rodents and not on that of North American or European hares and rabbits?

The third doubt was evoked by the fact that inhabitants of oceanic islands tended to resemble species found in the neighboring continents; African-like species in the Cape Verde Islands, South American-like species in the Galápagos archipelago. At the same time, although the geological and physical features of the Cape Verde and Galápagos Islands are very similar, their faunas are quite different. Why was this?

Finally, the different Galápagos islands, identical in climate and physical features, and very close to each other, might be expected to have identical species, but they had not. On different islands he observed that the finches characteristic of the Galápagos group differed in structure and food habits. Some ate insects, others ate seeds, and their sizes and their bills differed in relation to their diets and mode of life. Darwin suspected that they were only varieties of a single species. The local inhabitants could also tell at sight from which island any of the giant tortoises had come. Was all this arbitrary and meaningless or could a pattern of meaning be discerned?

The answer gradually came to Darwin: All these questions, and many more, could be rationally answered if species did not remain immutable but changed into other species, and diverged, so that one species could have given rise to two or more. It is because they had a common ancestor in South America that fossil armadillos resemble living armadillos, that agoutis resemble capybaras, and that the Galápagos birds resemble South American birds, while the Cape Verde birds resemble African birds with which in each case they had a common ancestor. But why did the finches and other birds of each Galápagos island differ from one another? “One might really fancy that, from an original paucity of birds in this archipelago, one species had been taken and modified for different ends,” Darwin wrote. These “ends” are very important; they fit the animals for their mode of life, they are adaptations; and the isolation of each species on each island plays an important part.

Gradually, only a few months after his return to England, these ideas began to crystallize in Darwin’s mind: “Animals, our fellow brethren in pain, disease, death, suffering and famine—our slaves in the most laborious works, our companions in our amusements—they may partake of our origin in one common ancestor—we may be all netted together.”

In July 1837 Darwin started writing down his ideas at random in his Notebook on Transmutation of Species. He soon found that if change of species had occurred, there was a ready explanation for a number of otherwise arbitrary and inexplicable facts. Why are the bones of the arm of a man, the foreleg of a dog and a horse, the wing of a bat, and the flipper of a seal built on the same general plan, with the different bones corresponding, each to each? Why are young embryos of lizards, chicks, and rabbits so similar to each other, while their adults are so distinct? Why do some animals have useless rudiments of organs? Why do particular regions of the earth have their characteristic plants and animals? Why do organisms fall into groups, namely, species, that may be arranged in larger groups, namely, genera, that in turn may be arranged in still larger groups, namely, families, and so on? Why is there this apparent relationship of species instead of a random distribution of forms across the whole field of possibilities? Why do more or less similar organisms behave in similar ways? For instance, why do both horses and men yawn; why do both orangutans and men show emotional distress by weeping? Why do fossils in early geological formations differ greatly from those living today, while those from more recent formations differ only slightly?

All these questions could be meaningfully answered if species were related by descent from common ancesters, but not otherwise. It is important to note Darwin’s form of argument. He never claimed to demonstrate the change of one species into another, and in his letters constantly repeated that he was tired of impressing this fact on his readers and critics; all that he claimed was that if evolution has occurred, it explains a host of otherwise inexplicable facts. But after he had satisfied himself that evolution had occurred, he kept his views to himself because he saw the great obstacles that lay ahead. Evolution he already saw as the change that species undergo in relation to their adaptation to their environment. A woodpecker differs from other birds in that it has two claws of its feet directed backward, with which it secures a firm grasp on the trunk of a tree, stiff tail feathers with which it supports itself against the tree, a stout bill with which it chisels a hole in the bark, and a very long tongue which it projects through the chiseled hole to extract the grubs beneath the bark. The woodpecker must have evolved these adaptations. How?

In considering this question of how evolution might have occurred, Darwin noticed that the cultivation of plants and the domestication and breeding of animals, both of which man has practiced since the Neolithic period, were the result of selection. Man deliberately chose the parents of the next generation of his plants and animals, to perpetuate and improve the qualities in them which he required. But how did, or could, selection operate in nature, where there had been no man to direct it, since the beginning of life on earth? Darwin had already observed that some species are better adapted than others to life in particular environments, and thus are likely to leave more descendants, while the less well adapted may diminish and become extinct (Notebook 1, MS. p. 37, notebook finished in February 1838). In other words, he had already grasped the principle of natural selection before he saw how it was enforced in nature.

The solution to this problem came to him on 28 September 1838 when he read Thomas Robert Malthus’ Essay on the Principle of Population. Malthus, whose whole line of thought was tinged with opposition to the principles of egalitarianism displayed by the French Revolution, argued that since the potential rate of increase in man was geometrical, and that his population could double in twenty-five years, while the increase of available foodstuffs could not increase so fast, there was bound to be misery, poverty, starvation, and death among the poor unless the rate of population increase in man was checked by war, famine, disease, or voluntary restraint (which would diminish the incentive to work among the poor).

Malthus, therefore, demonstrated the relentless pressure which human populations, by their boundless tendency to grow, maintain on their requirements for life whatever the level of their resources may be. If the food supply and other requirements for life can be increased, a population will tend to grow until its growth presses against the limits of the enlarged supply of food. Malthus also observed that in many countries the population did not grow and much of his book is taken up with a study of the means by which the numbers of people were limited in different countries. These ranged from exposure of infants, infanticide, and the intermittent reduction of the population by famine in China, to delayed marriage and strict sexual morality in Norway. But Malthus was able to show that in every country the tendency of a human population to grow was under some form of severe and continued restraint.

Darwin pounced on the argument and applied it, not to man, but to plants and animals in nature, and saw that they are in no position to increase their food supplies, but must die if they outstrip them by their own numbers. Here were the sanctions that made natural selection work.

Like most flashes of genius, it was very simple, and T. H. Huxley said later, “How extremely stupid not to have thought of that.” It marked a turning point, not only in the history of science, but in the history of ideas in general, for there is no field of human intellectual endeavor that has not been influenced by the thought and fact of evolution. By good fortune, the note that Darwin scribbled down immediately when the flash struck him has been preserved. It is of such importance that it deserves quotation:

28th [September 1838] We ought to be far from wondering of changes in numbers of species, from small changes in nature of locality. Even the energetic language of De Candolle does not convey the warring of species as inference from Malthus. Increase of brutes must be prevented solely by positive checks, excepting that famine may stop desire. In Nature production does not increase, whilst no check prevail, but the positive check of famine and consequently death. I do not doubt every one till he thinks deeply has assumed that increase of animals exactly proportionate to the number that can live.

Population is increase at geometrical ratio in FAR SHORTER time than 25 years, yet until the one sentence of Malthus no one clearly perceived the great check amongst men. There is [a] spring, like food used for other purposes as wheat for making brandy. Even a few years plenty makes population in man increase and an ordinun crop causes a dearth. Take Europe, on an average every species must have same number killed year with year by hawks, by cold etc. even one species of hawk decreasing in number must affect instantaneously all the rest. The final cause of all this wedging, must be to sort out proper structure, and adapt it to changes, to do that for form which Malthus shows is the final effect (by means however of volition) of this populousness on the energy of man. One may say there is a force like a hundred thousand wedges trying to force every kind of adapted structure into the gaps in the oeconomy of nature, or rather forming gaps by thrusting out weaker ones.

This passage, in which Darwin’s thought penetrates through a jungle of ideas, following on his reading of Candolle’s description of the war of nature and Malthus’ book, is remarkable for a number of reasons. In the first place it pinpoints the notion that evolution does not take place in a vacuum, but each individual, if it is lucky, lodges in its fortified position in the economy of nature, in what is now called its ecological niche, in dynamic equilibrium not only with the physical factors of the environment, but with the other living organisms of the habitat. It is because Darwin recognized this fact so clearly that he can be reckoned as the founder of the science of ecology.

Second, this passage shows that evolution is not a process that goes on in single individuals, or even in Paris, but which proceeds in populations and consists of some individuals becoming better adapted, under the pressure of natural selection, which is the third lesson of this passage. Ernst Mayr has pointed out that one of the most important recent advances in biology has been the realization that the real units are populations, which have objective existence, and not types (which are only imaginary abstractions), and that Darwin was more responsible than anyone else for the substitution of “population-thinking” for “typological thinking.” The change to a different species is nothing but a by-product of the process of a species becoming better adapted, after a certain point of divergence has been reached. The explanation of the fact of divergence itself is foreshadowed in the passage. As Darwin later wrote, “The more diversified the descendants of any one species become in structure, constitution and habits, by so much will they be better enabled to seize on many and widely diversified places in the polity of nature.” Again, it is the ecological niches which supply the key to the problem.

Darwin’s argument can be formulated as follows: (1) The numbers of individuals in species in nature remain more or less constant. (2) There is an enormous overproduction of pollen, seeds, eggs, larvae. (3) Therefore, there must be high mortality. (4) All individuals in a species are not identical, but show variation and differ from one another in innumerable anatomical, physiological, and behavioristic respects. (5) Therefore, some will be better adapted than others to their conditions of life and to the ecological niches which they could occupy, will survive more frequently in the competition for existence, will leave more off-spring, and will contribute most of the parents that will produce the next generation. (6) Hereditary resemblance between parents and offspring is a fact. (7) Therefore, successive generations will not only maintain but improve their degree of adaptation to their modes of life, i.e., to the conditions of their environments, and as these conditions vary in different places, successive generations will not only come to differ from their parents, but also from each other and give rise to divergent stocks issuing from common ancestors.

This is the formal theory of evolution by natural selection, which recent observation and controlled experiment have proved to be correct in all cases. What modern biology has done is to provide answers to the two questions which, in the state of knowledge of his time, Darwin could not answer: what is the nature of hereditary transmission between parents (and grandparents, and so forth) and offspring? And what is the nature of the origin of heritable variation? Without heritable variation, natural selection could achieve nothing. In Darwin’s day, no clear distinction was made between what was heritable and what was not, and few doubted the inheritance of characters acquired by a parent during its own life.

The problem was completely solved (unknown to Darwin, or to the world before 1900) by Gregor Mendel, who proved that the basis of inheritance takes the form of particulate characters, distributed between parents and offspring in accordance with a simple law (since proved by cytology and the behavior of chromosomes) known as Mendel’s law of segregation. The inception of a heritable novelty was a sport, now known as a mutation, a random, nonadaptive change in the chemical structure of a particular character, now known as a gene. The application of Mendelian genetics to Darwinian selection was effected by the work of J. B. S. Haldane and Sir Ronald Fisher by 1930. Since then the synthetic theory of evolution by natural selection has been generally accepted by biologists. It is remarkable that even without the knowledge that these advances represent, to say nothing of those achieved in comparative biochemistry, serology, parasitology, and cytogenetics, Darwin was nevertheless able to construct a coherent theory which made general acceptance of evolution possible—and a mechanism to account for it, namely, natural selection. Although long considered inadequate, natural selection is now proved to be the basis for biological change and for the production of all the diverse structures and functions of living organisms.

Shortly before Darwin died, his son Leonard asked him how long he thought that it would be before positive evidence of natural selection would be forthcoming. Darwin replied, about fifty years. It was a remarkably accurate forecast, for it was in the 1930’s that Sir Ronald Fisher showed how expression of a gene, in the form of the bodily character that it controls, is itself the effect of selection. Shortly afterward, E. B. Ford demonstrated that mimetic resemblance in butterflies is adaptive, confers survival value, and originates from heritable variation. The researches of H. B. D. Kettlewell on industrial melanism in moths, P. M. Sheppard and A. J. Cain in snails, T. Dobzhansky on geographical races of fruit flies, and A. C. Allison on sickle cell in man have provided experimental evidence not only of natural selection but of the pressure at which it works, and have permitted actuarial estimates of the longevity of different genetic types. This follows the mathematical studies of Sir Ronald Fisher, Sewall Wright, and J. B. S. Haldane on the effects of selection at different intensities on the composition of populations.

Studies on what B. Rensch has called “ring-races” has shown species in the act of splitting into new species, in gulls, tits, and salamanders, when geographical isolation of portions of populations enables them to evolve independently of each other, so that they become adapted in different directions and eventually no longer interbreed. This speciation is what T. H. Huxley regarded as necessary for the final confirmation of Darwin’s views.

Paleontology has been a trap for biologists and others who have neglected David Hume’s warning that the existence of a state of affairs does not in itself justify conclusions on how it came about.

Darwin claimed that the fossil record, as known to him, was compatible with evolution; but that if fossils ever provided any evidence contrary to his theory, such, for instance, as to prove that the Cambrian fossils were the first organisms that ever lived, or that a mammal was “created” later than those earliest known mammals of the Stonesfield Slate of the Jurassic period, his theory of evolution must be abandoned. However, he never adduced any evidence from fossils to bear on the problem of the principle of natural selection.

Although the recognition of fossils as former living beings goes back four centuries, it is only recently that paleontology has come into its own as a science, largely as a result of the researches of George G. Simpson. Fully aware that the mechanism of evolution can be interpreted only in terms of genetics, ecology, selection, and population studies, Simpson has shown that evolutionary sequences previously claimed as “straight” are not straight at all. For example, the evolutionary history of the horse has followed a zigzag course, first in the direction of many-toed browsers on leaves, then in that of many-toed grazers on grass, and finally in that of one-toed grazers on grass. In general the horse increased in size, but some species became smaller. Each lap of the course can be correlated with the environmental conditions of the period: soft or hard ground, leafy or grassy (siliceous) vegetation. At all times the trend has been toward favorable adaptations, to ecological niches which paleontologists are now in a position to describe; in other words, it has been compatible with the requirements of natural selection and has been opportunistic and devoid of any predetermined program. Furthermore, the rate of evolution, now measurable by radioactive dating, is not correlated with variability, nor with fertility (years per generation). In this manner, Simpson has been able to show that natural selection under changing environmental conditions accounts for evolution, and explains why in terrestrial forms evolution has generally been rapid, while in marine forms it has been slow or stationary.

It is regrettable that Darwin in later years allowed himself to be persuaded to accept Herbert Spencer’s inappropriate expression “survival of the fittest.” This stresses survival when the important thing is the greater fecundity of the better adapted; it emphasizes a superlative “fittest” when it is the slightest comparative superiority of adaptation that confers advantage. It lays the subject open to the taunt of tautology: Who survive? The fittest. Which are the fittest? Those who survive. This is a form of argument which neglects entirely the fundamental fact that forms better adapted to their environment leave more offspring. Finally, the phrase conveys no inkling that the automatic choice exerted by nature in ramming the better adapted variants into their ecological niches is the efficient cause of adaptation. It was not the first (or last) time that so-called philosophers of science have encumbered scientists with their help.

After Darwin had developed his new theory of the origin of species by natural selection in 1838, he did not publish it, or even discuss it with his friends. In 1842 he drew up a rough sketch of his argument, which in 1844 he expanded into an essay but never published. In 1845 when he had finished preparing for publication the results of his geological work on the Beagle voyage, he put the species question aside and started on eight tedious years’ study of the structure and classification of living and fossil barnacles. In the course of this work he acquired firsthand knowledge of the amount of variation that is found in nature, and he also made a striking discovery, that of complemental males, small parasitic males found under the mantle of larger hermaphrodite or female individuals.

After his return to England in 1836 Darwin became close friends with Charles Lyell and Joseph Dalton Hooker. They did not accept evolution, which was known to them only in the form of Lamarck’s exposition. Lamarck’s views were in many ways similar to those of Erasmus Darwin, who posited that there was a “natural tendency” to perfection, and the “inner a feelings” of an animal caused it to provide the organs required to meet its needs. In 1844 Robert Chambers published Vestiges of the Natural History of Creation, which only brought the subject into disrepute by its amateurish ignorance.

In April 1856 Darwin described his theory of natural selection to Lyell, who urged him to write a book describing his views on species. He began the work in the summer of 1856. On 18 June 1858 Darwin received a letter from Alfred Russel Wallace containing a perfect summary of the views which he had worked out during the preceding twenty years. Thanks to Lyell and to Hooker, Darwin’s and Wallace’s papers were read together before the Linnean Society of London on 1 July 1858 and published on 20 August of that year.

Darwin then set about writing an “abstract” of his larger work, and On the Origin of Species was published on 24 November 1859. The fat was then in the fire. Old-fashioned biologists protested that Darwin indulged in hypotheses that he could not prove (and he did not pretend to prove them). Theologians were aghast at two consequences of Darwin’s work; the first was that man and the apes must have had a common ancestor, which dislodged man from his privileged position as created by God in his own image. The second arose from the consideration that if Darwin’s views on the origins of plants and animals, including man, by natural selection were true, then much of the argument for the existence of God based on the presence of design in nature was destroyed. The theologian William Paley had argued that highly specialized and coordinated adaptations, such as the tongue, beak, claws, and tail of the woodpecker, or the human hand, manifested a complex design which required one to postulate the presence of a designer in nature. If, however, these same adaptations could be accounted for by natural selection acting on random variations, a designer was no longer needed. Since natural theology, as represented by Paley, had enjoyed enormous influence in England, the destruction of its very foundations was greeted with dismay and outrage.

Matters came to a head at the celebrated Oxford meeting of the British Association for the Advancement of Science on 30 June 1860, when the Reverend Samuel Wilberforce, bishop of Oxford (who knew little of natural history but was coached by the anatomist Richard Owen, who was jealous at what he felt already was Darwin’s ascendancy over himself), twitted Darwin’s friend Thomas Henry Huxley with the question whether it was through his father or his mother that he claimed descent from an ape. Huxley replied that he was not ashamed to be descended from an ape but he would be ashamed of an ancestor who used great gifts and eloquence in the service of falsehood. Wilberforce was annihilated by Huxley and Hooker, and Darwin’s views on evolution started their conquest of the world. In the United States, Darwin’s friend, the botanist Asa Gray, had already won a victory over the anatomist and paleontologist Louis Agassiz, who was never able to accept Darwin’s theory.

After the Origin of Species, Darwin wrote three more books expanding different aspects of the work. The Variation of Animals and Plants Under Domestication (1868) took up in detail that subject which had been confined to one chapter of the Origin. It contained his hypothesis of pangenesis, by means of which Darwin tried to frame an explanation of hereditary resemblance, inheritance of acquired characters, atavism, and regeneration. It was a brave attempt to account for a number of phenomena which were beyond the bounds of scientific knowledge in his day, such as fertilization by the union of sperm with egg, the mechanism of chromosomal inheritance, and the development of the embryo by successive cell division. His hypothesis of pangenesis could not therefore give a permanently acceptable account of the multitude of phenomena it was designed to explain. It was, however, a point of departure for particulate theories of inheritance in the later nineteenth century.

Darwin’s next book, The Descent of Man (1871), filled in what was only adumbrated in the Origin. T. H. Huxley and W. H. Flower had shown already that the body of man differs less from that of an ape than the latter does from that of a monkey. It is important to remember that Darwin never claimed that man was descended from apes, but that man’s ancestor, if alive today, would be classified among the Primates, and would be even lower in the scale than the apes. Man and apes are subject to similar psychological and physiological processes in courtship, reproduction, menstruation, gestation, birth, lactation, and childhood. At early stages of embryonic development the human fetus has a tail, inherited from ancestors who not only had a tail as embryos but also as adults and were quadrupedal. Darwin wrote, “The time will before long come when it will be thought wonderful, that naturalists, who were well acquainted with the comparative structure and development of man and other animals, should have believed that each was the work of a separate act of creation.”

Having proved his point with man’s body, Darwin then turned to the most difficult aspect, man’s mind. Here he showed that the gap, however enormous between man and the highest animal, is not unbridgeable, by the principle of gradation. Man shares with animals the urge to self-preservation, sexual love, maternal affection, paternal protection, and the senses of pleasure, pain, courage, pride, shame, excitement, boredom, wonder, curiosity, imitation, attention, and memory. As for the moral sense, Darwin concluded that it, also, arose gradually, through evolution, and that any animal endowed with social instincts, which man’s ancestors undoubtedly possessed, would acquire a moral sense as soon as its intellectual powers had developed (by natural selection) to an extent comparable with those of man and was able to appreciate the survival value of collaboration and mutual affection. Borrowing from Marcus Aurelius the view that social instincts are the prime principles of man’s moral constitution, Darwin concluded that these, with the aid of intellectual powers and the effects of new tradition and habits, would lead naturally to the Golden Rule: “As ye would that men should do to you, do ye to them likewise.”

In this book Darwin added an essay on a related subject: sexual selection, the preferential chances of mating that some individuals of one sex (usually the male) have over their rivals because of special structures, colors, and types of behavior used in courtship displays, leading them to leave more offspring and accentuate their characters: for example, antlers in deer, trains in peacocks, feathers in birds of paradise, color in sticklebacks. There is no need to assume a power of aesthetic choice in either sex; the characters act as sexual stimulants, real aphrodisiacs, so that mating follows more quickly, occurs more often, and there are more progeny. In man, there can be no doubt that the differences between the sexes in respect of distribution of hair on face and body and of fat in local accumulations under the skin are the result of sexual selection.

Darwin did not explain how the preference of females for particular structures, colors, and courtship displays was related to natural selection, although female preferences themselves must be subject to natural selection. Perhaps Darwin’s greatest contribution in this area was to show that secondary sexual characteristics had evolved in relation to a complex pattern of reproductive behavior, which must itself be the product of natural selection.

The last of Darwin’s books in this series, The Expression of the Emotions in Man and Animals (1872), contains studies of facial muscles and means of expression in man and mammals, emission of sounds, erection of hair, and so forth, and their correlation with suffering, sobbing, anxiety, grief, despair, joy, love, devotion, reflection, meditation, sulking, hatred, anger, pride, disdain, shame, surprise, fear, horror, acceptance (affirmation), and rejection (negation). With this book Darwin founded the study of ethology (animal behavior) and conveyance of information (communication theory) and made a major contribution to psychology.

Botanical Works. From the first Darwin interested himself in adaptations of plants to cross-pollination. He found that trees (with countless flowers) tend to have flowers of one sex, while small plants have hermaphrodite flowers, and he showed that the unisexuality of trees would tend to prevent self-pollination. He concluded that “flowers are adapted to be crossed, at least occasionally, by pollen from a different plant.” Here was a general principle which must have a wide meaning, and he set out to study it. His attention was first directed to orchids, which have elaborate adaptations for cross-pollination, making the bees that visit their flowers carry away the pollen sacs with them and pollinate the next flowers that they visit. His book on the Fertilization of Orchids (1862) showed that plants are in no way behind animals in the marvels of the adaptations that they show. Darwin further observed that flowers that are pollinated by wind have no colors; it is only those that are pollinated by insects that have bright colors in their petals and sweet-smelling nectaries.

Presently Darwin noticed that in some species flowers differ by the lengths of their anthers and styles, like the primroses, which show two conditions, or loosestrife, which shows three. This is also an adaptation for cross-pollination, and these observations formed the basis of Different Forms of Flowers on Plants of the Same Species (1877). The problem continued to fascinate him, and he raised two large beds of seedlings of Linaria vuglaris, the one cross-pollinated and the other self-pollinated, all from the same plant. “To my surprise, the crossed plants when fully grown were plainly taller and more vigorous than the self-fertilized ones.” Darwin had experimentally discovered and demonstrated the fact of hybrid vigor, or heterosis, which is completely explained by Mendelian genetics. These experiments, conducted over twelve years on fifty-seven species, led to the publication of Effects of Cross and Self Fertilization (1876). The demonstration of the advantage that accrues from cross-fertilization explains not only why sexual reproduction (as distinct from asexual budding) increases heritable variation (through recombination of genes), but also reveals the basis for the survival value conferred by the existence of different sexes in a species. This adaptation (for that is what it is) is a very old one, for it was inherited by plants and animals before they diverged from one another.

Darwin’s interest in climbing plants was not that of a systematic botanist but an attempt to discover what adaptive value the habit of climbing has. He found that “climbing” is a result of the process of nutation; the apex of the plant’s stem bends to one side while it grows and the plane of the bend itself revolves, clockwise or counterclockwise, so that the apex describes circular sweeping movements. In the hop plant—in hot weather, during daylight hours—it takes a little over two hours for each revolution. If the growing stem hits nothing, it continues to circle; if it hits an object it wraps itself around it by twining, Twining thus enables a young and feeble plant, in one season, to raise its growing point and leaves much higher from the ground, with more exposure to sunlight and air, without expending time and energy in the synthesis of woody supporting tissues. There is a further delicate adaptation here; a twining plant will not twine around an object larger than approximately six inches in diameter. This adaptation prevents it from climbing up a large tree, where it would be deprived of air and sun by the tree’s own leaves.

Some plants climb by a modification of this method and have stalks and leaves that clasp other objects, and intermediate forms show that leaf-climbers evolved from twiners. In others, such as the Virginia creeper, the leaf tendrils end in little discs full of resinous fluid which anchor the plant to its support. These researches were described in Climbing Plants (1875).

The behavior of climbing plants and the bending of their shoots led Darwin to investigate the mechanical cause of such bending. The fact that a stem bends toward the light because it grows faster on the unilluminated than on the illuminated side of the stem had been known for some time. By means of simple but ingenious experiments, Darwin showed that the tip of the shoot was sensitive to light and that the bending was caused by growth of the stem on the side away from the light but some way down from the apex. Furthermore, this growth was due to a substance that comes down from the apex, “some matter in the upper part which is acted upon by light, and which transmits its effects to the lower part.” From these researches and experiments (reported in Power of Movement in Plants, 1880) has sprung the whole science of growth hormones in plants.

Darwin’s chance observation of the number of flies caught on the leaf of the common sundew (Drosera rotundifolia) was the starting point for a series of observations and experiments which showed not only how insects are caught, but how their bodies are digested and ingested, and what the significance of this carnivorous habit is for the life of the plant. By feeding one bed of sundew plants with meat and depriving another bed, he found that the fed plants had larger leaves, taller flowerstalks, and more numerous seed capsules. This remarkable adaptation, which insures survival of the plants, explains why sundew plants, which have very few roots, through which only small supplies of nitrogen can be obtained, can live on extremely poor soil. Darwin was particularly impressed by the fact that the living cells of plants possess a capacity for irritability and response similar to that of the nerve and muscle cells of animals. His work on Insectivorous Plants was published in 1875.

Darwin’s last book connected with plants was The Formation of Vegetable Mould Through the Action of Worms (1881). This was published only six months before his death but covers a subject that he had studied for more than fifty years. He showed the services performed by earthworms in eating leaves and grinding earth in their gizzards and turning it into fertile soil, which they constantly sift and turn over down to a depth of twenty inches from the surface, thereby aerating it. He calculated from the weight of worm-castings that on one acre in one year’s time eighteen tons of soil are brought up to the surface by worms. This was a pioneer study in quantitative ecology.

Darwin was elected to fellowship of the Royal Society in 1839 at the age of twenty-nine; three foreign universities awarded him honorary doctorates; and fifty-seven leading foreign learned societies elected him to honorary or corresponding membership—the French Academy of Sciences only in 1878, for his views have never been really appreciated in France. The Prussian government awarded him the highly coveted Ordre pour le mérite, but from the British sovereign and the British government he never received any recognition. His demonstration of the fact of evolution, and of the automatic mechanism of natural selection which causes it, was unpalatable to the orthodox views of the Church of England. When he died, however, twenty members of Parliament asked the dean of Westminster to allow his burial in Westminster Abbey, an honor which was readily granted. Darwin himself would have been highly amused, for, with his genial sense of humor he once said, “Considering how fiercely I have been attacked by the orthodox, it seems ludicrous that I once intended to be a clergyman.” At his funeral there were present not only his friends Hooker, Huxley, and Wallace, but also James Russell Lowell, the American ambassador, as well as diplomats representing France, Germany, Italy, Spain, and Russia.

BIBLIOGRAPHY

I. Books. Journal of Researches into the Geology and Natural History of the Various Countries Visited by H. M. S. Beagle (London, 1839); facs. repr. (New York, 1952); 2nd ed., Journal of Researches into the Natural History and Geology of the Countries Visited During the Voyage of H. M. S. Beagle (London, 1845); repr., intro. by Sir Gavin de Beer (New York, 1956); The Structure and Distribution of Coral Reefs (London, 1842; 2nd ed., 1874); Geological Observations on the Volcanic Islands Visited During the Voyage of H. M. S. Beagle (London, 1844); Geological Observations on South America (London, 1846); A Monograph of the Subclass Cirripedia, 2 vols. (London, 1851, 1854); A Monograph of the Fossil Lepadidae, or Pedunculated Cirripedes of Great Britain (London, 1851); A Monograph of the Fossil Balanidae and Verrucidae (London, 1854); On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (London, 1859; 2nd ed., 1860; 3rd ed., 1861; 4th ed., 1866; 5th ed., 1869; 6th ed., 1872); repr., intro. by Sir Gavin de Beer (London, 1963); 1st American ed. (New York, 1860); variorum text, Morse Peckham, ed. (Philadelphia, 1959); facs. repr. of 1st ed. (Cambridge, Mass., 1964); On the Various Contrivances by Which British and Foreign Orchids are Fertilized by Insects, and on the Good Effects of Intercrossing (London, 1862, 2nd ed., 1877); The Variation of Animals and Plants Under Domestication (London, 1868; 2nd ed., 1875); The Descent of Man, and Selection in Relation to Sex (London, 1871; 2nd ed., 1874); The Expression of the Emotions in Man and Animals (London, 1872); Insectivorous Plants (London, 1875); Climbing Plants (London, 1875); The Effects of Cross and Self Fertilization in the Vegetable Kingdom (London, 1876); The Different Forms of Flowers on Plants of the Same Species (London, 1877); The Power of Movement in Plants, assisted by Francis Darwin (London, 1880); The Formation of Vegetable Mould, Through the Action of Worms, With Observations on Their Habits (London, 1881); repr., intro. by Sir Albert Howard (London, 1961); and a preliminary notice in Ernst Krause, Erasmus Darwin, W. S. Dallas, trans. (London, 1879).

Posthumously published writings include Life and Letters of Charles Darwin, Francis Darwin, ed. (London, 1887); Autobiography (London, 1887), unexpurgated ed., Nora Barlow, ed. (London, 1958); More Letters of Charles Darwin, Francis Darwin and A. C. Seward, eds. (London, 1903); “Sketch” of 1842 and “Essay” of 1844, in Evolution by Natural Selection, foreword by Sir Gavin de Beer (Cambridge, 1958); and the notebooks on transmutation of species, Sir Gavin de Beer, ed., in Bulletin of the British Museum (Natural History), Historical Ser., 2 (1960), 23–200, and 3 (1967), 129–176.

II. Bibliographies. Handlist of the Darwin Papers at the University Library Cambridge (Cambridge, 1960); R. B. Freeman, The Works of Charles Darwin. An Annotated Bibliographical Handlist (London, 1965).

III. Secondary Literature. See Sir Gavin de Beer, “Darwin’s Views on the Relations Between Embryology and Evolution,” in Journal of the Linnean Society of London, 66 (1958), 15–23; “The Origins of Darwin’s Ideas on Evolution and Natural Selection,” in Proceedings of the Royal Society, 155B , 321–338; “Mendel, Darwin, and Fisher,” in Notes and Records of the Royal Society, 19 (1964), 192–226; 21 (1966), 64–71; Atlas of Evolution (London, 1964); and Charles Darwin. A Scientific Biography (New York, 1965).

Also of interest are W. E. Le Gros Clark, Man-Apes or Ape-Men? (New York, 1967); A. Cronquist, The Evolution and Classification of Flowering Plants (London, 1967); T. Dobzhansky, Genetics and the Origin of Species (New York, 1937; 4th ed., 1959); Mankind Evolving (New Haven, 1962); R. A. Fisher, The Genetical Theory of Natural Selection (Oxford, 1930); E. B. Ford, Ecological Genetics (New York, 1964); Bentley Glass, Owsei Temkin, and William R. Strauss, eds., Forerunners of Darwin (Baltimore, 1959); Gerald L. Geison, “Darwin and Heredity: the Evolution of His Hypothesis of Pangenesis,” in Journal of the History of Medicine, 24 (1969), 375–411; Michael T. Ghiselin, The Triumph of the Darwinian Method (Berkeley, 1969); John C. Greene, The Death of Adam. Evolution and Its Impact on Western Thought (Ames, Iowa, 1959); J. B. S. Haldane, The Causes of Evolution (London, 1932); Garett Hardin, Nature and Man’s Fate (London, 1960); Julian Huxley, A. C. Hardy, and E. B. Ford, eds., Evolution as a Process (London, 1954); Julian Huxley, Evolution. The Modern Synthesis (London, 1942; repr., 1963); Ernst Mayr, Animal Species and Evolution (Cambridge, Mass., 1963); Milton Millhauser, Just Before Darwin. Robert Chambers and Vestiges (Middletown, Conn., 1959); Bernard Rensch, Evolution above the Species Level (London, 1959); and Anne Roe and George Gaylord Simpson, eds., Behavior and Evolution (New Haven, 1958).

See also George Gaylord Simpson, The Meaning of Evolution. A Study of the History of Life and of its Significance for Man (London, 1950); Horses. The Story of the Horse Family in the Modern World and Through Sixty Million Years of History (New york, 1951); The Major Features of Evolution (New York, 1953); This View of Life. The World of an Evolutionist (New York, 1964); The Geography of Evolution (Philadelphia-New York, 1965); G. Ledyard Stebbins, Variation and Evolution in Plants (New York, 1957); and Sol Tax, ed., Evolution After Darwin, 3 vols (Chicago, 1960).

On Darwin’s health see Saul Adler, “Darwin’s Illness,” in Nature, 184 (1959), 1102–1103.

Gavin de Beer

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"Darwin, Charles Robert." Complete Dictionary of Scientific Biography. 2008. Encyclopedia.com. 26 May. 2016 <http://www.encyclopedia.com>.

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Darwin, Charles

Darwin, Charles

WORKS BY DARWIN

SUPPLEMENTARY BIBLIOGRAPHY

Charles Robert Darwin was born in 1809 at Shrewsbury and died in 1882 in Down House in Kent. He was the next to youngest child in a family of four sisters and an older brother. His father, Robert Darwin, was a physician of imposing presence and severe character. His paternal grandfather, Erasmus, had also been a doctor by profession. By avocation, however, Erasmus Darwin was a naturalist, whose Zoönomia, a disquisition in verse on the course of nature, has usually been considered, quite incorrectly, to have been an anticipation of Charles Darwin’s theory of evolution. Darwin’s maternal grandfather was Josiah Wedgwood, founder of the pottery firm and friend of Erasmus Darwin in the Birmingham scientific circle known as the Lunar Society. Darwin belonged, in short, to a provincial family of means and intelligence which associated itself with the professions, intellectual pursuits, and commerce. Although his father expected him to choose a profession, he never needed to earn a living. In 1839 Darwin, too, married a Wedgwood, his cousin Emma. The seven (of ten) children who survived infancy and their descendants have sustained the notable literary and scientific tradition which distinguishes the family.

Originally intended for medicine in his turn, Darwin detested the study of it at the University of Edinburgh. His father allowed him to move to Cambridge, with a view to his becoming a clergyman. At both universities the hobby of natural history absorbed him. He read; he collected beetles and prepared specimens; he went on geological and entomological forays. He might have become a clerical naturalist like Gilbert White, whose Natural History and Antiquities of Selborne had charmed him in boyhood. Instead, a decisive opportunity intervened. In 1831 he was offered the (unpaid) post of naturalist aboard H. M. S. Beagle, then outfitting for a voyage to collect geographical information. This was to be no mere trip around the world. The expedition lasted five years. It was the making of Darwin as a scientist, and it cost him his health. Sir Gavin de Beer’s biography has established the strong probability that the chronic physical prostration which debilitated Darwin throughout the rest of his life was the manifestation of Chagas’ disease, a malady identified only in 1909, which involves infestation of the heart muscle and intestines by Trypanosoma cruzi. These bacteria are carried by the great black bug of the pampas, benchuca (Triatoma infestans), and while in Peru Darwin recorded having watched with careful interest one of these insects biting his finger and engorging itself on his blood.

Darwin returned to England in 1836. On the Origin of Species appeared in 1859. In the interval of over twenty years Darwin published descriptive accounts of the zoological and geological findings of the voyage. His Journal of Researches in 1839 was followed by a series of geological treatises: one on coral reefs in 1842, a second on volcanic islands in 1844, and a third on South America in 1846. These writings won him the reputation of an accurate, thorough, and reliable naturalist. In the early 1850s he published in four volumes a complete study of fossil and living barnacles. Here his work was that of a descriptive biologist of great finesse, capable also in experimental inquiry. Meanwhile, he was quietly maturing his views on evolution, these too deriving from observations and reflections started on the voyage of the Beagle.

Theory in biology. The theoretical originality of Darwin’s central work is more difficult to appreciate than its content is difficult to understand. On the Origin of Species does not on the face of it read like a book animated by theory, either abstruse or abstract. Paragraphs follow one out of another setting forth evidence for the mutability of animals. That notion was not in itself a new one. Intimations and even evidence of the transformation of species had often before been adduced—by Lamarck, by Darwin’s grandfather, and by many others. Certainly Darwin marshaled his materials with a fidelity to fact and a range of relevance that proved telling. Nevertheless, what was original was not the assertion but the explanation of evolution in the theory of natural selection. For that theory constituted a new notion of biological order. Although Darwin’s mundane style tended to obscure its novelty, the Origin is in fact a work of theory; it changed the way of seeing the phenomena of life.

Prior to the assimilation of Darwin’s work, no consensus existed about the mode of biological explanation. Only in the early nineteenth century did biology as a professional discipline emerge from the combination of the tradition of natural history with that of the medical or mechanistic study of anatomy and physiology. In Darwin’s youth two distinct traditions governed the modeling of theory—the one organismic and the other teleological. The former arose in eighteenth-century romantic feeling about nature and harked back to ancient Stoicism. According to that philosophy, nature is the source of vitality, morality, and destiny. Goethe’s vision of universal metamorphosis exemplified this cast of mind, and its foremost scientific exponent was Lamarck. In post-Darwinian polemics Lamarck has often been represented by those hostile to the theory of natural selection as the predecessor who had formulated whatever was original and valid in the doctrine of evolution. That argument is untrue to the facts and unfair to the thought of both Lamarck and Darwin. In Lamarck’s view, cyclic equilibrium obtains in the world between the continuously constitutive forms of activity (electricity, fire, life itself) and the perpetually disintegrating tendency of the physical environment. The innate self-perfecting drive of living beings never altogether prevails over the dead hand of inorganic matter, and the tableau of animals each in its kind represents the highest point to which every species has attained in the continuous process of becoming. Assertions of the inheritance of acquired characters and the law of use and disuse (these are later terms) may, indeed, be found in Lamarck’s writings, although only as incidental to his view of flux and process in the progression of life, not as explaining actual evidence of variations.

Over against this conception of life as plastic power, novel in emphasis and appeal if not in principle, persisted the strong tradition of teleology. Engrained since Aristotle, that habit of mind had survived the scientific revolution of the seventeenth century with its scope reduced to the degree that it had become the mode of explanation appropriate to the description of organisms. Within this restricted domain teleology had if anything been fortified by the popularity in eighteenth-century natural theology of the argument from design, which held that the existence of God is demonstrable in the ease and skill with which He had specially created each species to fit its own environment and manner of life. In scientific practice, moreover, teleology answered in detail to the problems of functional analysis of anatomical structure, exhibiting how in plants and animals the part serves in exact respects the purposes of the whole. Such was Cuvier’s method in comparative anatomy.

Background of “Origin of Species.” The problem of adaptation, in short, was the crux of biological explanation. Aboard the Beagle Darwin already saw it to be so. Leaving home a believer in the fixity of species, like Cuvier or Paley, he had the facts of geographical variation of species continuously borne in upon him throughout the extensive experience of the voyage. He told in his Autobiography what lines of evidence led him first to doubt and then to change his mind:

… I had been deeply impressed by discovering in the Pampean formation great fossil animals covered with armour like that on the existing armadillos; secondly, by the manner in which closely allied animals replace one another in proceeding southwards over the [South American] Continent; and thirdly, by the South American character of most of the productions of the Galapagos archipelago, and more especially by the manner in which they differ slightly on each island of the group. ([1876a] 1958, p. 118)

Upon these manifold indications of variation of species in geographical space the new science of geology superimposed evidence of their modification in time. Stratigraphy had become a definitive science in the decades between 1810 and 1830. The first truly historical science, stratigraphy systematically ordered the geological succession according to the fossil population characteristic of the several formations and systems of the rocks. While an undergraduate, Darwin had worked one summer as assistant to one of the founders of stratigraphy, the Reverend Adam Sedgwick, a professor at Cambridge. Sedgwick, like many among the early geologists, held that the evidence for geological revolutions confirmed Biblical accounts of the Creation and the Flood. That view proved venial scientifically, however, for their intensive study of the fossil record made clear the progression of forms of life in the course of the ages.

Catastrophists of Sedgwick’s persuasion took those ages to be brief and punctuated by cataclysms. Increasingly, however, skeptical critics came to think of the span of the past as very long—long enough for all changes to come about gradually. Indeed, the mentor to whom Darwin professed the deepest of his scientific obligations was not a biologist at all. He was Sir Charles Lyell, whose Principles of Geology, published in three volumes between 1830 and 1833, was the earliest fully comprehensive work in that science. It combated the doctrine of geological catastrophes in the name of the uniformity and virtual eternity of nature and taught Darwin to think of the cumulative efficacy of minute changes operating progressively across vast reaches of time.

Further insights came to Darwin as he puzzled over the problem of species in the months following his return to England in the autumn of 1836. Soon thereafter he started a notebook on the subject, the first in what was to become a voluminous series. He worked the while, so he later said, on Baconian principles, collecting facts wholesale. One of his concerns was inheritable modifications. Such modifications are not to be confused with characteristics acquired, in the Neo-Lamarckian sense, as a consequence of a creature’s individual necessities—the biceps in a blacksmith’s arm, for example. Any theory of evolution must, of course, suppose that the variations which are its subject matter are cumulative over generations. From the outset Darwin had the wit to see that if a theory of evolution were not to beg the question, variations must be supposed random in origin, cumulative in effect, and indefinite in extent. In thinking about how they may be accumulated, Darwin meditated often upon the way in which kennel owners, cattle farmers, stablemen, nurserymen, and pigeon fanciers select stock for breeding—how, operating upon successive generations of domesticated animals and plants, the breeder picks for reproduction those individuals that exhibit the characters desired and thus appears to mold the species.

As early as 1837, then, Darwin had all this: geographical evidence of variation; geological proof of succession; a uniformitarian sense of the history of nature; and the notion of random, heritable, and cumulative modifications in form, modifications produced by a process of selection, albeit an artificial one. Even so, he saw no prospect of oversetting by means of such indirect evidence the well-nigh universal conviction that species are fixed in nature itself, based as that belief was on the universal observation that within the experience of everyone like reproduces like, as well as on the plausible supposition that the adaptation of species to all sorts of specialized lives argues the creation of each to lead just that life we see it living.

Darwin disposed of a set of facts and ideas, in other words, and needed a theory; indeed, he was constantly looking for one. It came in one of those flashes of insight which is better called catalytic than coincidental or subconscious. In October 1838 he chanced to read for distraction Maithus’ essay on population. The argument persuaded him at once [seePopulation, article onpopulation theories; and the biography ofalthus]. It must, indeed, have been familiar, for it was one of the commonplaces of contemporary political economy, requiring only to be brought together with Darwin’s problem for its relevance to flash into his mind:

… being well prepared to appreciate the struggle for existence which everywhere goes on from long-continued observation of the habits of animals and plants, it at once struck me that under these circumstances favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species. Here, then, I had at last got a theory by which to work. ([1876a] 1958, p. 120)

He worked for twenty years before publishing, although he did write out a draft in 1844 for circulation among friends and colleagues, notably Lyell and J. D. Hooker, the botanist. In May 1856 he at last began writing on an exhaustive scale and completed some ten or eleven chapters in the next year and a half. He thought to call the work “Natural Selection.” Then on June 18, 1858, Darwin had the shock of receiving from Alfred Russel Wallace, a young naturalist who had been investigating the flora and fauna of the Malayan jungle, the manuscript of an evolutionary theory identical with his own although based on much less extensive material, together with a request to send it along to Lyell if it should seem worthy. This coincidence precipitated Darwin’s decision to come before the public. On the advice of Lyell and Hooker he prepared a brief summary of his views to be published jointly with Wallace’s paper. Thereupon he settled down to compose an account that should be full enough to do justice to his materials although reduced in scale from what he had initially intended. It took eight months to write the Origin of Species, which appeared in November 1859.

The circumstances of the publication of the Origin belong, of course, to Darwin’s personal history, not to an account of his science. Suffice it to say that his behavior throughout was entirely honorable. It offers no grounds for the imputation repeated by detractors that he slighted Wallace. (For a general discussion of multiple discoveries in science, see Merton 1961.)

“Origin of Species.” The full title summarizes the argument: On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. Addressing himself to the general public, Darwin began with the most familiar matter, the great variations evident among domesticated animals. He thereby called attention to the variability which exists within what are commonly taken to be fixed species and observed that in the breeder’s hands they are malleable: “The key is man’s power of accumulative selection: nature gives successive variations; man adds them up in certain directions useful to him” ([1859] 1964, p. 105). The second chapter extends consideration of the ambiguity of species into nature, and the third attributes the function of selection to the struggle for existence. “Owing to this struggle, variations, however slight, and from whatever cause proceeding, if they be in any degree profitable to the individuals of a species, in their infinitely complex relations to other organic beings and to their physical conditions of life, will tend to the preservation of such individuals, and will generally be inherited by the offspring” (p. 145). Within the species the proportion of favored individuals will increase in successive generations, since by definition these individuals have characteristics with survival value that enable their possessors, merely by living longer than less favored competitors, to produce more descendants similarly equipped. Thus will species alter as time goes on.

The middle chapters of the Origin discuss the laws of variation. Darwin’s masterful control over detail is particularly evident here. There are acute observations on the effects of use and disuse of organs, on the influence of climate, and on the greater variability of species than of genera, the last offering a consideration peculiarly difficult to comprehend on any basis other than his. Darwin expressly excluded the problem of cause. Given the evidence that variations have in fact occurred and are heritable, the theory of natural selection explains the course of variation or change. Indeed, the success of the theory turned on dropping the question of cause, not as uninteresting, but as currently unanswerable, although not necessarily forever so.

In the later chapters Darwin anticipated difficulties and objections. He dealt with instinct, which hardly seems a bodily matter. He acknowledged the problem of extreme specialization and discussed the spectacular beauty of the male in certain species of birds as an extravagance that seems to surpass the humdrum effects of selection. He ascribed the absence of transitional forms to the imperfection of the geological record. He treated sexual preferences and choices as a special case of natural selection rather than as an auxiliary or extraneous factor. In summary let it be said that contemporary evolutionists bear him out on all essential matters. The degree of confirmation of his reasoning and its fidelity to vast reaches of fact is extraordinary when it is also considered that he was necessarily unaware of the twentieth-century discoveries about the fine structure and workings of heredity.

Publication of the Origin was the great event, indeed the raison d’être, of Darwin’s career. His single-mindedness invested the book with a kind of power—a power deriving from the scope as well as from the depth of his thought. Iconoclastic critics have sometimes described the book as dull. Nor does it invite stylistic appreciation. It will repay structural analysis, however, and this is the way to understand Darwin’s merit as a theorist. For this is a work of theory, even though any given sentence expresses only fact or deduction. The art lies in the deployment, not in the expression, and it creates its effect in the way a mosaic does rather than a painting. Each element is a mere bit of stone, every sentence a mere statement; yet a design governs the arrangement and the disposition. There are not many books in just this genre. Burckhardt’s Civilization of the Renaissance in Italy, for example, and perhaps Marx’s Capital (setting aside the disgust which often overlays statements in Marx) are also works in the mid-nineteenth-century mode of factual argumentation which, in the literature of science, Darwin represents at its most powerful.

Darwin did not abandon further science in order to defend his views in the discussion that followed the publication of the Origin, although the pattern of the remaining half of his career was inevitably determined by the response to his book. He followed criticism closely and modified later editions of the Origin in numerous respects which either met or accepted objections without abandoning the position. Not all the revisions proved fortunate. Least felicitous was one he introduced into the subtitle as well as into the text. At Wallace’s urging he substituted Herbert Spencer’s formula, “survival of the fittest,” for his own phrase, “preservation of favoured races” to describe what transpires in the struggle for life in consequence of natural selection. Both the tautology—i.e., survival in a struggle for life—and the ambiguity of the superlative “fittest” confused his proper meaning and obscured the point he had originally emphasized: that the essential feature in the operation of natural selection is not mere survival for its own sake. It is the differential reproductivity of individuals favored adaptively by chance variation, which effect entails an increasing proportion of similarly constituted offspring in a population.

Darwin’s later work. Thenceforth, Darwin was more effective in his other writings, both in substantiation of the theory of natural selection and in affirmation of his remarkable virtuosity in research. He chose problems which combined an appeal to his taste with relevance to evolution. Three major books develop central evolutionary themes more fully than does the Origin. The Variation of Plants and Animals Under Domestication (1868) tells in full zoological and archeological detail how man has bent to his uses very variable breeds—dog and horse, for example, sheep and goat, peacock and guinea fowl—and attributes the contrasting stability of the goose to its never having been taken in as a pet or servant and of the donkey to its having served only the classes of society whose members cannot afford experiments [seeDomestication, article onAnimal domestication]. By way of excursion, at the end of his book Darwin advanced the hypothesis of pangenesis to account for the phenomena of heredity. Here he addressed himself to the complaint that he had not explained how variations are transmitted. The speculation with which he met the criticism has only increased his vulnerability before later critics, for in imagining that the basis of heredity might be generalized throughout the organism (“gemmules” in each structure and organ governing the formation of its counterpart in the offspring), Darwin appeared to have conceded inheritance of acquired characters taken in a Lamarckian sense. It is to be appreciated, however, that Darwin suggested this speculation as a parenthetical aside.

In The Descent of Man (1871) Darwin directed his theory explicitly to the single species Homo sapiens. Much of the book is devoted to developing the evolutionary significance of sexual selection— i.e., the preferential choice of reproductive partner. Apart from that, the argument follows two main lines. The first is physical. The immediate evolutionary forebears of mankind are unknown. Darwin never represented man as deriving from apes. He did establish that man must in all probability be descended from species that are classified among primates, and further, that man and the higher apes resemble each other anatomically more closely than the latter resemble the lower primates.

The second line of argument is behavioristic. Intellectual and social faculties are themselves adaptive and in their variations make for the greater or lesser survival of the creatures that possess them. Other species besides man subsist with the aid of rudimentary or developed forms of social organization and communication, and Darwin could see only differences of degree between these and the characteristics of human community and moral awareness.

The last of Darwin’s expressly evolutionary treatises carries behavioristic comparisons even further. Probably The Expression of the Emotions in Man and Animals (1872) is the most dated of Darwin’s writings. Nevertheless, in comparing phenomena like the physical manifestation of hostility in dog and master—the similarity of the snarling jaw to the drawn lip—and many other states of temper, Darwin did carry biology into treatment of faculties traditionally reserved for moral studies. Reciprocally, he introduced the study of animal behavior into psychology [seeSocial behavior, animal].

In the last decade of his life Darwin put in hand a train of experimental researches mainly botanical in character and published a final series of monographs exhibiting a great variety of novel observations and phenomena. He had already brought out a study of fertilization in orchids—in 1862; in 1877 he issued a second edition, much enlarged in detail. In the same year he published a very acute work on hermaphroditic reproduction in trees and plants. Two years previously, in 1875, he had issued two quite new studies, one on climbing plants (first published in journal form in 1865) and one on insectivorous plants. In 1880 he generalized the former in a book about the power of movement in plants as it resides in root, leaf, stem, and flower. In 1881 he concluded his lifework with an account of the process by which earthworms contribute to the formation of humus: the originality of the subject is characteristic of his acumen, while its choice reflects the modesty of his temperament.

To appreciate the full range of Darwin’s accomplishment as a scientist one must read in these experimental writings as well as in the Origin. The comparison to Newton is often adduced. It is justified. Notable scientists normally excel either in theoretical insight or in experimental virtuosity. Newton and Darwin were capable as very few have ever been, and perhaps none in equal measure, of combining those veins and thereby achieving the foremost stature in all aspects that distinguish science. It is true that, given the state of the science, comprehensiveness had to do the work in theory for Darwin that mathematical power did for Newton, which is only to say that the one was suited by his abilities to accomplish for biology what the other had done for physics, and that both of them made the most of those abilities.

Professional response to the “Origin.” The theory of natural selection polarized opinion among biologists more radically than is the common effect of scientific theories. Those who adopted it tended to do so straightaway in something like a conversion. “How extremely stupid not to have thought of that,” was T. H. Huxley’s often quoted reaction as the pieces suddenly fell into place on reading Darwin. Yet even the professional response was influenced by philosophical taste and cultural tradition. Then, as now, the English-speaking scientific world was empirically minded after the manner of Bacon, Locke, and Mill. In Darwin’s own milieu, what immediately persuaded was his resolution of the question of biological adaptation. Not that he solved the problem. He did better. He dispelled it, by turning adaptation from final cause into present effect. Instead of a wonder so intricate as to bespeak Providence or purpose, adaptation became merely the outcome of the whole life history of the species in its relation to circumstance and environment, a phenomenon or set of phenomena to be analyzed and explained in evolutionary terms. In Huxley’s hands, therefore, Darwin became the protagonist of the drama of scientific fact undoing theological prejudice.

In Germany, where Darwin’s work enjoyed a success if anything more influential than in England and the United States, what appealed was less its empiricism than its historicism. The sense of history as indwelling process, whether in the successive versions of Herder, Hegel, or Marx, had in the course of the romantic movement of thought become deeply engrained in the German sense of the world. Philosophically speaking, such historiography treats the past in a mode exactly counter to Darwin’s, the one being a realist and the other a nominalist approach to nature. Oblivious to these distinctions, German historicists misread Darwin and took him for their own. It is well known that Marx spoke of dedicating the second book of Capital to Darwin, an intention which makes a curious contrast to the Lamarckian position of some subsequent Marxists. The professional champion and self-appointed spokesman, however, was Ernst Haeckel, who tried to make over all biology into the study of evolution. He governed his embryology, for example, by the doctrine that the development of every individual in the womb recapitulates the evolution of the species. Not only does evolution govern all biology, but all truth too: Haeckel’s Riddle of the Universe is the cardinal document of the movement to make something like a religion out of evolutionary thought itself.

Only in France was Darwin’s work initially greeted by indifference (except among anthropologists who did take it up with very great benefit to their science). Not so the biologists. Among them the writ of Cuvier still ran. There was also a problem of language. In French, the word évolution refers to the development of an individual; transformisme connotes the doctrine as affecting species in general, and at that time involved visionary connotations. A self-defeating semantic objection also supervened—to explain the origin of species in a book which questioned their reality seemed an inadmissible mode of discourse. The greatest of physiologists, Claude Bernard, never even seized the differences between the Darwinian theory and its speculative predecessors. “We must doubtless admire,” he wrote disdainfully, “those great horizons dimly seen by the genius of a Goethe, an Oken, a Carus, a Geoffrey Saint-Hilaire, a Darwin, in which a general conception shows us all living beings as the expression of types ceaselessly transformed in the evolution of organisms and species, —types in which every living being individually disappears like a reflection of the whole to which it belongs” ([1865] 1957, pp. 91–92). In general, Darwin expressed himself pragmatically in a manner characteristically English and characteristically unconvincing, not to say irritating, to the French.

The nonprofessional response. Darwin himself took no part in the social Darwinist movement of ideas and gave it no countenance [see Social Darwinism]. Something must here be said, nevertheless, about the pattern of nonprofessional response, insofar as it has been hostile, for these attitudes greatly amplified for a time certain scientific difficulties left unresolved by Darwin, and they have also contributed to persistent misunderstandings of the status of the theory in science—misunderstandings arising from repugnance toward its political and social derivations.

The pattern has two distinct aspects, theological and moralistic, which turn on different implications of the theory of evolution. The former, the theological issue between the Biblical story of the Creation and the Darwinian account of the origin of species, is the more famous. It is also the more literal and has proved the more ephemeral, concerned as it is with the wide historical fact of evolution rather than with the theory of natural selection, by which Darwin explained and thereby established the fact. The science of geology had, indeed, already faced theology with the necessity of regarding the scriptural account of earth history in an allegorical fashion, and the confrontation between Darwinists and religious believers settled rather than initiated that matter among informed persons. The Origin of Species did cause additional shock and distress, of course, chiefly in regard to the question of whether the reality of a spiritual component of human existence requires that man be biologically unique and specially created. This is a dilemma which may be resolved in one of two ways: either by considering the whole universe as contingent and composed of physical and spiritual aspects; or, in a more restricted way, by appreciating that the theory of evolution treats man as an evolved animal, but it does not and in principle cannot assert that he is nothing but an animal. At all events, within a relatively short space of time theologians learned to contemplate evolution without dismay, as clearly a matter of fact; significant religious minds have not felt impelled to gird against the facts, although fundamentalist rejection of them persists in diminishing backwaters.

What has persisted in recurrent and often serious manifestations is a second aspect of resistance to Darwin’s work. This kind of resistance is serious because it obtains among educated persons and reveals the occasional inability of highly intelligent people to respect the force and limitations of scientific demonstration. In this tradition the tenor has been to accept and even celebrate the phenomenon of evolution while denigrating the theory of natural selection. Sometimes this denigration has taken the form of an attack on Darwin’s character and probity, as in the instances of Barzun (1941) and Himmelfarb (1959). Sometimes it has taken the form of a revival, more or less garbled, of the views of Lamarck, as in the writings of Samuel Butler and George Bernard Shaw. Sometimes it has taken the form of proposing an alternative formula, metaphorical in character, for the purpose of replenishing evolution with intelligence, will, or choice. Such were the notions of “creative evolution,” the elan vital of Bergson, or entelechy of Driesch. Sometimes it has taken on ideological and political dress, as in the refusal of Lysenko’s adherents to admit the irrelevance to heredity of manipulation of the physical and social environment. As, indeed, in any complex of opinions, some proportion of these several elements, some mixture of these motivations, may be discerned in all types of educated refusal to be persuaded by the theory of natural selection.

What unifies these hostile views into a tradition of romantic resistance is the common complaint that the theory of natural selection deprives the history of nature of any sanctions for man as a social or moral being. These critics hold a different conception of the function of a scientific explanation. They want more out of nature than Darwin, or science generally, has found there. They want reasons, and Darwin gave them circumstance. They want causes, and he gave them accidental variation. They want purpose and direction, and he gave them an indefinite sequence of indifferent results. They want a science which seizes on unity and deep process, and he fragmented nature into a congeries of discrete events connected only by proximity and happenstance. What has proved profound, therefore, is not the incompatibility between evolutionary science and theology, considered intellectually at least. Much deeper and more persistent, partly because of common commitment to the natural with different understandings of what it signifies, has been the incompatibility between the Darwinian theory and any form of naturalistic social or moral philosophy. For the moralist knows what sort of nature he requires of science. And the theory of natural selection has succeeded precisely in consequence of limiting its purview to the indifferent event. While the present state of scientific understanding is not the touchstone by which to judge theories of the past, it is important to emphasize, nevertheless, that the theory of natural selection is part of working evolutionary thought. To refuse to admit its force is not only to question the actual state of scientific knowledge, but it is also to impugn the current mode of scientific explanation.

Charles C. Gillispie

[Directly related are the entriesEvolution; Genetics; Race. Other relevant material may be found in the biographies ofCooley; Galton; Giddings; Lubbock; Morgan, Lewis Henry; Spencer; Tylor.]

WORKS BY DARWIN

(1839) 1952 Journal of Researches Into the Geology and Natural History of the Various Countries Visited by H.M.S. Beagle. New York: Hafner. → A facsimile reprint of the first edition. A second edition was published in 1845.

(1842) 1962 The Structure and Distribution of Coral Reefs. Berkeley: Univ. of California Press. → A second edition was published in 1874.

(1842–1858) 1958 Darwin, Charles R.; and Wallace, Alfred R. Evolution by Natural Selection. With a foreword by Gavin de Beer. Cambridge Univ. Press. → The Darwin and Wallace papers of 1858, together with Darwin’s other early drafts.

(1844–1846) 1900 Geological Observations on the Volcanic Islands and Parts of South America Visited During the Voyage of H.M.S. “Beagle.” 3d ed. New York: Appleton. → First published as Geological Observations on the Volcanic Islands, Visited During the Voyage of H.M.S. Beagle (1844) and Geological Observations of South America (1846).

1851 A Monograph on the Fossil Lepadidae: Or, Pedunculated Cirripedes of Great Britain. London: Palaeontological Society.

1851–1854 A Monograph on the Sub-class Cirripedia, With Figures of All the Species. 2 vols. London: Ray Society. → Volume 1: The Lepadidae: or, Pedunculated Cirripedes. Volume 2: The Balanidae (or Sessile Cirripedes), the Verrucidae, etc.

1854 A Monograph on the Fossil Balanidae and Verrucidae of Great Britain. London: Palaeontological Society.

(1859) 1964 On the Origin of Species. Cambridge, Mass.: Harvard Univ. Press. → A facsimile of the first edition. Numerous editions have been published, including a reprint of the sixth edition in 1927 by Macmillan, and a variorum text edited by Morse Peckham, published by the Univ. of Pennsylvania Press in 1959.

(1862) 1903 The Various Contrivances by Which Orchids Are Fertilized by Insects. 2d ed., rev. New York: Appleton. → First published as On the Various Contrivances by Which British and Foreign Orchids Are Fertilised by Insects.

(1865) 1893 The Movements and Habits of Climbing Plants. 2d ed., rev. New York: Appleton. → First published in Volume 9 of the Journal of the Linnean Society.

(1868) 1900 The Variation of Animals and Plants Under Domestication. New York: Appleton.

(1871) 1930 The Descent of Man and Selection in Relation to Sex. 2d ed., rev. enl. New York: Appleton.

(1872) 1965 The Expression of the Emotions in Man and Animals. Edited by Francis Darwin. Univ. of Chicago Press.

(1875) 1900 Insectivorous Plants. 2d ed., rev. Edited by Francis Darwin. New York: Appleton.

(1876a) 1958 Autobiography of Charles Darwin, 1809–1882. Edited by his granddaughter Nora Barlow. London: Collins. → Written in 1876; first published posthumously in 1887.

(1876b) 1902 The Effects of Cross and Self Fertilisation in the Vegetable Kingdom. New York: Appleton.

(1877) 1896 The Different Forms of Flowers on Plants of the Same Species. New York: Appleton.

1880 The Power of Movement in Plants. London: Murray.

(1881) 1892 The Formation of Vegetable Mould Through the Action of Worms, With Observations on Their Habits. New York: Appleton.

(1887) 1959 The Life and Letters of Charles Darwin. 2 vols. New York: Basic Books. → Includes an autobiographical chapter, edited by Francis Darwin.

SUPPLEMENTARY BIBLIOGRAPHY

Barzun, Jacques (1941) 1958 Darwin, Marx, and Wagner: Critique of a Heritage. 2d ed. Garden City, N.Y.: Doubleday.

Bernard, Claude (1865) 1957 An Introduction to the Study of Experimental Medicine. New York: Dover. → First published in French.

De Beer, Gavin R. 1964 Charles Darwin: Evolution by Natural Selection. New York: Doubleday. → The most authoritative biography.

Eiseley, Loren 1959 Darwin’s Century: Evolution and the Men Who Discovered It. Garden City, N.Y.: Doubleday.

Gillispie, Charles C. 1951 Genesis and Geology: A Study in the Relations of Scientific Thought, Natural Theology, and Social Opinion in Great Britain, 1790–1850. Harvard Historical Studies, Vol. 58. Cambridge, Mass.: Harvard Univ. Press. → A paperback edition was published in 1959 by Harper.

Glass, Bentley (editor) 1959 Forerunners of Darwin: 1745–1859. Baltimore: Johns Hopkins Press.

Himmelfarb, Gertrude 1959 Darwin and the Darwinian Revolution. Garden City, N.Y.: Doubleday.

Merton, Robert K. 1961 Singletons and Multiples in Scientific Discovery: A Chapter in the Sociology of Science. American Philosophical Society, Proceedings 105:470–486.

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Darwin, Charles Robert

DARWIN, CHARLES ROBERT

(b. The Mount, Shrewsbury, England, 12 February 1809;

d. Down House, Downe, Kent, England, 19 April 1882),natural history, geology, evolution. For the original article on Darwin see DSB, vol. 3 (note error in date of birth given there).

When Gavin de Beer wrote the article on Darwin in the original DSB there were only a handful of historical studies available, including his own biography. Over the next few decades there was an explosion of interest in the process by which Darwin developed his theory of evolution by natural selection, fueled by the publication of his notebooks, edited by de Beer himself. There have also been many studies of the impact of Darwin’s ideas, inspired in part by a recognition that earlier perceptions had been shaped by the success of the selection theory in the mid-twentieth century. Michael Ruse coined the term “Darwin industry” to denote the group of historians who devoted themselves to the study of his work. Many of Darwin’s papers have since been published, including a combined volume of the notebooks originally edited by de Beer. There are editions of his diaries, of the marginalia he scribbled in the books he read, and an ongoing project to publish his whole correspondence. Both the Origin of Species and the notebooks have been provided with a concordance. The flow of technical literature has slackened off, but the flood of biographical studies has continued unabated, those by Janet Browne and by Adrian Desmond and James Moore having attracted particular attention.

There have been important developments in the understanding of many aspects of Darwin’s work and its implications. Historians such as Jonathan Hodge have alerted scholars to the strong thread that runs through Darwin’s work based on a traditional view of heredity and reproduction. There have been major debates on the role of ideology in his thinking and new developments in the understanding of his changing religious views. De Beer’s effort to trace a straightforward link between Darwin’s original theory and modern Darwinism have come under suspicion as researchers have become more aware of the existence of alternative ideas about how evolution might work.

Darwin’s Early Career . Darwin’s period of medical study at Edinburgh has traditionally been dismissed as of little significance. But he also worked on natural history there, and this has led Hodge and Phillip R. Sloan to challenge the impression given in his autobiography that these were wasted years. Darwin claimed to have been unimpressed by the Lamarckian anatomist Robert Grant, although it is now known that the two worked closely together, and that Darwin was impressed by Grant’s claim that the “zoophytes” (Hydrozoa and corals) serve as a bridge between the plant and animal kingdoms. Darwin’s early reading focused his attention on issues that would shape his later thinking, especially his ideas about generation or sexual reproduction. He became committed to the view that reproduction is a creative activity of the vital forces in the body, a position quite unlike the modern focus on the rigid transmission of genetic units. Here his thinking reflects the fascination with the creativity of sex that ran through his grandfather Erasmus Darwin’s theories.

Sandra Herbert has made a major study of Darwin’s geological work and has shown how he turned the methodology of Charles Lyell into a theoretical program for understanding all Earth movements. There have been important studies of Darwin’s biogeographical work on the Beagle voyage. Working on Darwin’s original notes, Frank Sulloway has undermined the myth of a “eureka” experience on the Galápagos Islands by showing that the finches there could not have played such a crucial role. Darwin’s own specimens were not even labeled to show which island they were collected on, and he had to rely on collections made by others on the ship to reconstruct the distribution. Only after his return to England, when the ornithologist John Gould informed him that the finches really did constitute a group of distinct but closely related species, did Darwin realize their full significance.

Herbert and Martin Rudwick have made detailed studies of Darwin’s active role in the London geological community following his return, including his abortive theory on the parallel roads of Glen Roy. But behind the scenes he had begun to think hard about the question of transmutation. Sulloway argues that the insights that led Darwin to the idea of branching evolution were probably arrived at after the Beagle had returned to England in October 1836, although a few historians speculate that he may have accepted transmutation during the later part of the voyage. Ernst Mayr called this the first Darwinian revolution: The Galápagos results convinced him that new species are formed by the natural transformation of old ones, although as yet he had no idea what the process of change might be. Over the next few years he searched for a plausible mechanism, and his development of the theory of natural selection constitutes a second revolution.

The Discovery of Natural Selection . Howard Gruber has explored Darwin’s notebooks from this period to argue that the discovery of the selection theory can be seen as a major example of creative thinking. Darwin sometimes projected an image of himself as a patient observer, and his opponents have always accused him of being incapable of deep thought. But he also said that the Origin was “one long argument,” and Mayr among others has stressed that he was anything but a simple fact-gatherer. Ruse and Silvan Schweber focus on Darwin’s desire to appear a good scientist as defined by contemporary discussions of the scientific method by Sir John Frederick William Herschel and William Whewell. They argue that he developed natural selection by a creative process of synthesis and testing. David J. Depew and Bruce H. Weber see the selection theory as the product of a new statistical mode of explanation and also stress his innovative work in the area of methodology. Darwin wanted to create a theory based on natural law in the Newtonian tradition, but by the very nature of the problems he addressed he was forced to transform this program by introducing statistical and historical elements into his explanations. Robert J. Richards sees the historical element as derived from German idealist philosophy, but others, including Hodge, relate it to Lyell’s uniformitarian methodology.

Darwin started from his knowledge of biogeography, and with a conviction that evolution must be a branching process in which one species is divided by geographical barriers and the separate populations then become transformed in different directions. He wondered if species might be “born” with a built-in lifespan, after which they become extinct. He soon realized, however, that some more active mechanism of change was needed and by July 1837 he was convinced that transmutation must come about by the accumulation of individual variations over many generations. He explored the direction already taken by Jean-Baptiste de Lamarck and Erasmus Darwin: Might a change in the environment produce modifications either by affecting the reproductive process or by changing the organisms’ habits? He soon decided that Lamarckism was inadequate. For a variety of reasons, he decided that although the environment might well be the stimulus, the majority of the changes it produced were not purposeful.

Studies by Hodge, David Kohn, and Phillip Sloan have revealed the extent to which Darwin was influenced by his views on “generation” or reproduction. The key to transmutation was the creative power of sexual reproduction. The theory of heredity he later called “pangenesis” (published in 1868) was formulated at this early stage. Older studies lament Darwin’s “failure” to anticipate genetics, on the assumption that if he had developed the concept of the gene he would have avoided the problems that beset the selection theory in the late nineteenth century. Modern scholars accept that the foundations of his thought lie in a premodern concept of heredity.

James Secord and Peter J. Vorzimmer show how the C notebook shows Darwin studying work of animal breeders, where changes could actually be observed within a species. He later suggested that he observed the breeders applying a process of artificial selection and then saw how this model could be transferred to natural evolution. L. T. (Lloyd Thomas) Evans and Ruse have explored the analogy between artificial and natural selection because—as Darwin himself later realized—it offers such a neat model for how natural selection operates. But although the process of selection seems clearly illustrated in such cases, Darwin’s notebooks show that there was no direct borrowing of the selection model from the breeders. Throughout the C and D notebooks he remained convinced that adaptive variations must somehow be elicited automatically in a changed environment (Lamarckism), and did not see the breeders work as a useful analogy. Accounts of the discovery of the selection mechanism by Camille Limoges, Herbert, and Richard A. Richards have argued that he could not have been inspired by the model of artificial selection. Darwin became so used to citing the work of animal breeders as a model in his later accounts of the theory that he actually came to believe that this was how he was led to the idea of selection.

At this point he read Thomas Robert Malthus on population and realized the significance of the struggle for existence. Here the discussion enters the debate over the extent to which his thinking reflected the ideology of free-enterprise capitalism. Robert M. Young has been the most powerful advocate of the view that the selection theory does reflect the prevailing ideology of the time, a position backed up in the biography by Desmond and Moore. Schweber argues for an influence derived from the individualism of Adam Smith’s economics, while noting that scientific factors had already begun to make Darwin think in terms of individual variation within a population—the innovation that Mayr calls “population thinking”—the transition from seeing a species as based on an ideal type to seeing it as a population of distinct individuals. Depew and Weber see the rise of Darwinism as the creation of a statistical mode of explanation as opposed to the old Newtonian view of causation based on law. The claim for a direct input from Darwin’s social environment must be balanced against the evidence for a growing awareness that for science to tackle certain kinds of questions, a new type of explanation based on statistically modeled changes was needed.

De Beer’s account minimized the influence of Malthus, treating the population principle as only a catalyst helping Darwin to put together insights already gained from his observations of nature. The connection is certainly not as direct as is sometimes implied: Peter J. Bowler showed that Darwin’s concept of the struggle for existence does not appear in Malthus’s work, and in this sense he had to think creatively with the insight provided by the population principle. Malthus did not anticipate the logic of what became known as social Darwinism, and thus could not provide the whole model for the selection theory. Edward Manier and Schweber point to the breadth of Darwin’s reading, which reveals other ways in which the ideology of the time could have filtered into his thinking. He read Smith and also David Brewster’s review of Auguste Comte’s positivist philosophy, which argued for the need to base all theories on mathematical foundations—exactly what was provided by the arithmetical logic of the population principle. Looking for a way of measuring variation, he turned to the work of the Belgian anthropologist Lambert Quételet, who pioneered the application of statistics to the human population. Darwin’s theory was not a slavish copy of Malthus’s political philosophy, but it may nevertheless have been steeped in the ideology of the time. As Desmond and Moore stress, this was a time of social conflict in Britain, so Darwin could see on the streets the struggle that the political economists were trying to understand.

Studies by Gruber and Herbert show the M and N notebooks revealing Darwin’s adoption of a materialist perspective, at least on how his theory would apply to humankind. These notes anticipate many of the topics later articulated in the Descent of Man. Darwin saw no room for the traditional notion of a soul existing on a purely spiritual plane: The mind was a product of the material activity of the brain, just as the phrenologists held. He suspected that much of human unconscious behavior may be instinctive, programmed into human brains by the effect of evolution on humanity’s ancestors. The ways in which individuals express their emotions reveal their animal ancestry, as in the case of snarling to express anger. He was convinced that evolution would throw light on moral values by showing how certain forms of social behavior have been programmed into humans by natural selection. Morality was merely a rationalization of these social instincts. Such ideas would undermine the whole traditional view of human nature.

Yet Darwin still felt that the laws of nature were instituted by a wise and perhaps even benevolent God. Far from recognizing the full horror of a worldview based on struggle, he stressed that the end result of evolution was to keep species well adapted to their environments in an ever-changing world. As Walter Cannon and Young argue, he transformed rather than destroyed the old natural

theology of William Paley (1743–1805). Even struggle and death had a positive value in the divine plan. Robert Richards argues that Darwin was also committed to the idea of progress. He suggests that historians influenced by modern Darwinism have failed to appreciate that Darwin still saw evolution as a steady pressure to mount the scale of organization. Selection was a force that would tend to raise the standard of organization whenever the circumstances were appropriate. But he was also aware that the advance could take place in a number of directions, not just along a single line leading toward humanity.

Development of the Theory . Over the next twenty years Darwin worked on his theory, soon moving to live in the countryside at Downe. He was not a complete recluse, however, and the vast extent of the correspondence network he built up (later published) provided him with sources of information on subjects relevant to his theorizing. As Manier argues, he was trying to build up a community of scientists who would speak the new language of evolution.

Darwin’s work on barnacles provided him with many insights, including the importance of embryological characters for determining relationships. His thinking incorporated an element of what would later be called the recapitulation theory, although the extent to which he can be associated with the full-blown version of this theory is controversial. Richards has argued that Darwin adopted the “law of parallelism,” whose influence had been charted by Stephen Jay Gould. This form of recapitulation theory implies a linear model of evolution, with the lower animals being treated as immature versions of the more perfect human form. Many historians are suspicious of Richards’s effort to tie Darwin in with the developmental way of thinking so prevalent in Germany.

An important change took place in Darwin’s thinking in the 1850s. It has often been assumed that as soon as he formulated the selection theory, he must have recognized that it implied a much harsher vision of nature as a scene of unrelenting struggle and suffering. But Dov Ospovat argued that the early form of the theory still revealed its roots in Paley’s vision of natural theology. Only in the 1850s did he realize that a compromise could not be maintained. The principle of population implies that the struggle for existence continues even in a stable environment, so there can never be any escape from the relentless threat of death. As Frank Burch Brown, Kohn, and Neal Gillespie show, Darwin’s worldview thus became gradually more pessimistic. Moore notes how this move was encouraged by the death of his beloved daughter Anne in 1851.

Ospovat also notes that this growing recognition of the power of struggle allowed Darwin to explain the trends toward divergence and specialization being discovered by Richard Owen in the fossil record. Both Owen and William B. Carpenter recognized this trend by comparing the fossil sequences with the process of embryological specialization observed by Karl Ernst von Baer. In effect, Darwin applied the principle of the “division of labor” to explain why it was advantageous for species to diversify. Browne and Kohn note that biogeography also transformed Darwin’s thinking on this issue. His “principle of divergence” arose from a recognition that successful genera expand and diversify, wiping out the species that had previously occupied the territory. Here again Darwin came face to face with the relentless pressure of natural selection. Sulloway notes that by focusing on the pressure to specialize, Darwin was led to marginalize his early emphasis on the role of geographical isolation in speciation, as in the Galápagos. Nevertheless, by 1856 he had created a much more sophisticated theory and had amassed a vast amount of supportive evidence and had begun to write a “big book” on the topic, now published and edited by Robert Stauffer. This was the project interrupted by the arrival of Alfred Russel Wallace’s paper in 1858. There has been much debate over the degree of similarity between the two men’s theories of natural selection, summarized by Malcolm Kottler. Some scholars argue that there were significant differences between them, but no one disputes that Darwin recognized important parallels and began writing the Origin of Species.

The Reception of Darwin’s Theory . Surveying the recent work on the reception of the Origin of Species moves into broader realms of the interactions between Darwin and the scientific, cultural, and social developments of the time. A survey of the Darwinian revolution by Ruse provides a useful overview, as do the biographies of Darwin by Browne and Desmond and Moore. Recent studies of the early Darwinians such as Thomas Henry Huxley should also be consulted.

For the scientific debate, David Hull edited and commented on a collection of reviews of the Origin. Scholars such as Vorzimmer, Jean Gayon, and William Provine have traced the debates over natural selection from Darwin’s time into the era of modern genetics. The emergence of the so-called modern synthesis of Darwinism and genetics was the topic of an important collection of essays edited by Mayr and Provine. Here is a complex series of debates centered on challenges to the traditional assumption that Darwin’s “failure” to recognize the significance of Gregor Mendel’s laws of heredity left him vulnerable to attacks such as the one launched by Fleeming Jenkin in 1867. In the early 2000s, recognition of the role played by a pregenetical model of heredity and variation in Darwin’s thinking about selection makes it difficult to imagine how he could have appreciated the potential value of Mendelism. Nor is it clear that his theory of pangenesis necessarily undermined the credibility of the selection mechanism, although Jenkin’s attack did force him to question his longstanding assumption that favorable variations were very rare. The process by which the theory of natural selection was synthesized with genetics was long and complex because it involved the destruction of a developmental viewpoint in which the transmission of characters was thought to be inextricably connected with the mechanism by which those characters are generated in the embryo.

Many modern historians would accept that it was the prevalence of this developmental model in late nineteenth-century biology, not a specific problem associated with “blending inheritance” that accounts for the widespread reluctance to take the theory of natural selection seriously during Darwin’s own lifetime. Peter Bowler has outlined several alternative theories of evolution developed in the late nineteenth-century “eclipse of Darwinism.” In particular he develops Gould’s account of the ways in which the recapitulation theory was used to imply that evolution must be directed along predefined channels, just like the development of the embryo. Bowler has also suggested that an appreciation of the extent to which Darwin’s own explanation of the process was rejected must force us to reconsider the reasons why the basic idea of evolution became popular at the time. Darwinism was translated into a theory of necessary progress, and the more radical implications of the selection theory remained dormant until the twentieth century. Along similar lines, Ruse has written a provocative book arguing that Darwinism succeeded only because the theory of evolution piggybacked on the rising tide of enthusiasm for the idea of progress.

In terms of the wider public response, Alvar Ellegard provided a detailed study of the reaction to Darwin in the British periodical press. Gillian Beer has pioneered the study of Darwin’s influence in the literary sphere. Moore has challenged the established view that evolutionism became sucked into a “war” between science and religion, noting the extent to which liberal Christians welcomed the theory because they saw it as a model for social progress in the modern world. The challenges posed by this vision to the more traditional Christian view of the origin of humanity were highlighted in Darwin’s own study of human origins, the Descent of Man. Richards has written a detailed account of the development of evolutionary explanations of the human mind, which compares Darwin’s ideas on the topic with those of other thinkers such as Herbert Spencer. He stresses the extent to which Darwin and Spencer saw themselves as providing morality with a new foundation, certainly not as destroying all traditional values. Darwin did, however, come to accept the prevailing view that the nonwhite races retain more traces of humanity’s ape ancestry, although initially he had adopted more liberal views on this issue.

Curiously, however, there have been comparatively few attempts to integrate Darwin’s ideas on human origins with the history of paleoanthropology, although a survey by Bowler stresses the innovative nature of his belief that the development of an upright posture was the key breakthrough in the separation of the first hominids from the apes. Virtually all other commentators before the mid-twentieth century took it for granted that it was the expansion of the human brain that drove the separation— Darwin was the first to suggest that the initial step might have been triggered by an adaptive change necessitated by the hominids’ move onto the open plains.

There have been endless debates about the role played by Darwin’s theory in the promotion of “social Darwinism.” Historians such as Young, who stress the ideological input into Darwin’s own thinking, have naturally pointed to the use of Darwinian rhetoric to justify policies in which struggle is the driving force of progress. There have been many accounts of social Darwinism making the same point, from the classic study by Richard Hofstadter to more recent work by Mike Hawkins. But historians such as Robert Bannister and Bowler have urged caution, noting that some appeals to the metaphor of the struggle for existence did not depend on the application of the selection theory, even when Darwin’s name was invoked. Spencer’s enthusiasm for individual competition was linked to a Lamarckian mechanism of self-improvement, for all that he coined the term “survival of the fittest.”

There have been comparatively few accounts of the detailed biological investigations Darwin carried out in the later part of his life, apart from those contained in the biographies. Helena Cronin and Kottler have written on the theory of sexual selection, especially Darwin’s longstanding debate with Wallace on the topic (significantly in the context of the “eclipse” of Darwinism; few other biologists took that theory seriously until the mid-twentieth century). On a more personal level, Ralph Colp has written on Darwin’s illness. Moore has discredited the story, popular among modern creationists, that he underwent a deathbed conversion back to Christianity. Moore has also provided an analysis of the symbolism associated with the events surrounding his funeral. When he died in 1882 the scientific community was anxious to exploit his status as the most famous biologist of the century to obtain for him the honor of being buried in Westminster Abbey.

SUPPLEMENTARY BIBLIOGRAPHY

There have been many modern editions of The Origin of Species, usually based on either the first or the sixth editions, and translations into most major languages.

WORKS BY DARWIN

Charles Darwin’s Natural Selection: Being the Second Part of His Big Species Book Written from 1856 to 1858. Edited by Robert C. Stauffer. London: Cambridge University Press, 1975.

The Collected Papers of Charles Darwin. Edited by Paul H. Barrett. 2 vols. Chicago: University of Chicago Press, 1977.

A Concordance to Darwin’s Origin of Species, First Edition. Edited by Paul H. Barrett, Donald J. Weinshank, and Timothy T. Gottleber. Ithaca, NY: Cornell University Press, 1981.

The Correspondence of Charles Darwin. Edited by Frederick Burkhardt and Sydney Smith. Cambridge: Cambridge University Press, 1985–. Thirteen volumes published thus far, series still in progress.

A Calendar of the Correspondence of Charles Darwin, 1821–1882. Edited by Frederick Burkhardt, Sydney Smith, David Kohn, et al. New York and London: Garland, 1985.

Charles Darwin’s Notebooks, 1836–1844: Geology, Transmutation of Species, Metaphysical Enquiries. Edited by Paul H. Barrett, Peter Jack Gautrey, Sandra Herbert, et al. London: British Museum (Natural History); Ithaca, NY: Cornell University Press, 1987.

Charles Darwin’s Beagle Diary. Edited by Richard D. Keynes. Cambridge, U.K.: Cambridge University Press, 1988.

Charles Darwin’s Marginalia. Vol. 1. Edited by Mario di Gregorio and Nick Gill. New York: Garland, 1990–.

A Concordance to Charles Darwin’s Notebooks, 1836–1844. Edited by Donald J. Weinshank et al. Ithaca, NY: Cornell University Press, 1990. Available from www.darwin-online.org.uk

OTHER SOURCES

Bannister, Robert C. Social Darwinism: Science and Myth in Anglo-American Social Thought. Philadelphia: Temple University Press, 1979.

Beer, Gillian. Darwin’s Plots: Evolutionary Narrative in Darwin, George Eliot, and Nineteenth-Century Fiction. London: Routledge and Kegan Paul, 1983.

Bowler, Peter J. “Malthus, Darwin, and the Concept of Struggle.” Journal of the History of Ideas 37 (1976): 631–650.

———. The Eclipse of Darwinism: Anti-Darwinian Evolution Theories in the Decades around 1900. Baltimore: Johns Hopkins University Press, 1983.

———. Evolution: The History of an Idea. Berkeley: University of California Press, 1984. Rev. ed., 2003.

———. Theories of Human Evolution: A Century of Debate, 1844–1944. Baltimore: Johns Hopkins University Press, 1986.

———. The Non-Darwinian Revolution: Reinterpreting a Historical Myth. Baltimore: Johns Hopkins University Press, 1988.

———. Charles Darwin: The Man and His Influence. Oxford: Basil Blackwell, 1990.

Brown, Frank Burch. “The Evolution of Darwin’s Theism.” Journal of the History of Biology 19 (1986): 1–45.

Browne, Janet. “Darwin’s Botanical Arithmetic and the ‘Principle of Divergence,’ 1854–1858.” Journal of the History of Biology13 (1980): 53–89.

———. Charles Darwin: A Biography. 2 vols. London: Cape, 1995–2002.

Cannon, Walter F. “The Bases of Darwin’s Achievement: A Reevaluation.” Victorian Studies5 (1961): 109–134.

Colp, Ralph. To Be an Invalid: The Illness of Charles Darwin. Chicago: University of Chicago Press, 1977.

Cronin, Helena. The Ant and the Peacock: Altruism and Sexual Selection from Darwin to Today. Cambridge, U.K.: Cambridge University Press, 1991.

Crook, Paul. Darwinism, War, and History: The Debate over the Biology of War from the “Origin of Species” to the First World War. Cambridge, U.K.: Cambridge University Press, 1994.

Depew, David J., and Bruce H. Weber. Darwinism Evolving: Systems Dynamics and the Genealogy of Natural Selection. Cambridge, MA: MIT Press, 1995.

Desmond, Adrian, and James R. Moore. Darwin. London: Michael Joseph; New York: Viking Penguin, 1991.

Ellegard, Alvar. Darwin and the General Reader: The Reception of Darwin’s Theory of Evolution in the British Periodical Press, 1859–1872. Göteburg, Sweden: Acta Universitatis Gothenburgensis, 1958. Reprint, Chicago: University of Chicago Press, 1990.

Evans, L[loyd] T[homas]. “Darwin’s Use of the Analogy between Artificial and Natural Selection.” Journal of the History of Biology 17 (1984): 113–140.

Gayon, Jean. Darwinism’s Struggle for Survival: Heredity and the Hypothesis of Natural Selection. Cambridge, U.K.: Cambridge University Press, 1998.

Gillespie, Neal C. Charles Darwin and the Problem of Creation. Chicago: University of Chicago Press, 1979.

Gould, Stephen Jay. Ontogeny and Phylogeny. Cambridge, MA: Harvard University Press, 1977.

Gruber, Howard E. Darwin on Man: A Psychological Study of Scientific Creativity, by Howard E. Gruber, Together with Darwin’s Early and Unpublished Notebooks, Transcribed and Annotated by Paul H. Barrett. New York: E. P. Dutton, 1974.

Hawkins, Mike. Social Darwinism in European and American Thought, 1860–1945: Nature as Model and Nature as Threat. Cambridge, U.K.: Cambridge University Press, 1997.

Herbert, Sandra. “The Place of Man in the Development of Darwin’s Theory of Transmutation.” Pts. 1 and 2. Journal of the History of Biology7 (1974): 217–258; 10 (1977): 155–227.

———. Charles Darwin, Geologist. Ithaca, NY: Cornell University Press, 2005.

Hodge, Michael J. S. “The Structure and Strategy of Darwin’s ‘Long Argument.’” British Journal for the History of Science 10 (1977): 237–246.

———. “Darwin and the Laws of the Animate Part of the Terrestrial System (1835–1837): On the Lyellian Origins of His Zoonomical Explanatory Program.” Studies in the History of Biology 6 (1983): 1–106.

———. “Darwin as a Lifelong Generation Theorist.” In The Darwinian Heritage, edited by David Kohn. Princeton, NJ: Princeton University Press, 1985.

———, and David Kohn. “The Immediate Origins of Natural Selection.” In The Darwinian Heritage, edited by David Kohn. Princeton, NJ: Princeton University Press, 1985.

———. “Generation and the Origin of Species (1837–1937): A Historiographical Suggestion.” British Journal for the History of Science 22 (1989): 267–282.

Hofstadter, Richard. Social Darwinism in American Thought. Rev. ed. New York: George Braziller, 1959.

Hull, David L. Darwin and His Critics: The Reception of Darwin’s Theory of Evolution by the Scientific Community. Cambridge, MA: Harvard University Press, 1973.

Kohn, David. “Theories to Work By: Rejected Theories, Reproduction, and Darwin’s Path to Natural Selection.” Studies in the History of Biology 4 (1980): 67–170.

———. “Darwin’s Principle of Divergence as Internal Dialogue.” In The Darwinian Heritage, edited by David Kohn. Princeton, NJ: Princeton University Press, 1985.

———. “Darwin’s Ambiguity: The Secularization of Biological Meaning.” British Journal of the History of Science 22 (1989): 215–240.

———, ed. The Darwinian Heritage. Princeton, NJ: Princeton University Press, 1985.

Kottler, Malcolm Jay. “Darwin, Wallace, and the Origin of Sexual Dimorphism.” Proceedings of the American Philosophical Society124 (1980): 203–226.

———. “Charles Darwin and Alfred Russel Wallace: Two Decades of Debate over Natural Selection.” In The Darwinian Heritage, edited by David Kohn. Princeton, NJ: Princeton University Press, 1985.

Limoges, Camille. La sélection naturelle: Étude sur le première constitution d’un concept (1837–1859). Paris: Presses Universitaires de France, 1970.

Manier, Edward. The Young Darwin and His Cultural Circle: A Study of Influences which Helped Shape the Language and Logic of the First Drafts of the Theory of Natural Selection. Dordrecht: Reidel, 1978.

———. “History, Philosophy, and Sociology of Biology: A Family Romance.” Studies in the History and Philosophy of Science11 (1980): 1–24.

Mayr, Ernst. “Darwin and Natural Selection: How Darwin May Have Discovered His Highly Unconventional Theory.” American Scientist65 (1977): 321–327.

———. The Growth of Biological Thought: Diversity, Evolution, and Inheritance. Cambridge, MA: Belknap Press, 1982.

———. One Long Argument: Charles Darwin and the Genesis of Modern Evolutionary Thought. Cambridge, MA: Harvard University Press, 1991.

———, and William B. Provine, eds. The Evolutionary Synthesis: Perspectives on the Unification of Biology. Cambridge, MA: Harvard University Press, 1980.

Moore, James R. The Post-Darwinian Controversies: A Study of the Protestant Struggle to Come to Terms with Darwin in Great Britain and America, 1870–1900. New York: Cambridge University Press, 1979.

———. “Charles Darwin Lies in Westminster Abbey.” Biological Journal of the Linnean Society17 (1982): 97–113.

———. The Darwin Legend. Grand Rapids, MI: Baker, 1994.

Ospovat, Dov. “The Influence of Karl Ernst von Baer’s Embryology, 1828–1859: A Reappraisal in Light of Richard Owen and William B. Carpenter’s ‘Paleontological Application of von Baer’s Law.’” Journal of the History of Biology 9 (1976): 1–28.

———. The Development of Darwin’s Theory: Natural History, Natural Theology, and Natural Selection, 1838–1859. Cambridge, U.K., and New York: Cambridge University Press, 1981.

Provine, William B. The Origins of Theoretical Population Genetics. Chicago: University of Chicago Press, 1971.

Richards, Richard A. “Darwin and the Inefficiency of Artificial Selection.” Studies in the History and Philosophy of Science 28 (1997): 75–97.

Richards, Robert J. Darwin and the Emergence of Evolutionary Theories of Mind and Behavior. Chicago: University of Chicago Press, 1987.

———. The Meaning of Evolution: The Morphological Construction and Ideological Reconstruction of Darwin’s Theory. Chicago: University of Chicago Press, 1992.

Rudwick, Martin J. S. “Darwin and Glen Roy: A ‘Great Failure’ in Scientific Method?” Studies in the History and Philosophy of Science 5 (1974): 97–185.

———. “Charles Darwin in London: The Integration of Public and Private Science.” Isis73 (1982): 186–206.

Ruse, Michael. “The Darwin Industry: A Critical Evaluation.” History of Science 12 (1974): 43–58.

———. “Charles Darwin and Artificial Selection.” Journal of the History of Ideas36 (1975): 339–350.

———. “Darwin’s Debt to Philosophy: An Examination of the Influence of the Philosophical Ideas of John F. W. Herschel and William Whewell on the Development of Charles Darwin’s Theory of Evolution.” Studies in the History and Philosophy of Science6 (1975): 159–181.

———. The Darwinian Revolution: Science Red in Tooth and Claw. Chicago: University of Chicago Press, 1979.

———. Monad to Man: The Concept of Progress in Evolutionary Biology. Cambridge, MA: Harvard University Press, 1996.

Schweber, Silvan S. “The Young Darwin.” Journal of the History of Biology12 (1979): 175–192.

———. “Darwin and the Political Economists: Divergence of Character.” Journal of the History of Biology13 (1980): 195–289.

———. “John Herschel and Charles Darwin: A Study in Parallel Lives.” Journal of the History of Biology 22 (1989): 1–71.

Secord, James A. “Nature’s Fancy: Charles Darwin and the Breeding of Pigeons.” Isis72 (1981): 163–186.

Sloan, Phillip R. “Darwin’s Invertebrate Program, 1826–1836.” In The Darwinian Heritage, edited by David Kohn. Princeton, NJ: Princeton University Press, 1985.

———. “Darwin, Vital Matter, and the Transformism of Species.” Journal of the History of Biology 19 (1986): 369–445.

Sulloway, Frank J. “Geographic Isolation in Darwin’s Thinking: The Vicissitudes of a Crucial Idea.” Studies in the History of Biology 3 (1979): 23–65.

———. “Darwin and His Finches: The Evolution of a Legend.” Journal of the History of Biology15 (1982): 1–54.

———. “Darwin’s Conversion: The Beagle Voyage and Its Aftermath.” Journal of the History of Biology15 (1982): 325–396.

Vorzimmer, Peter J. Charles Darwin: The Years of Controversy; The Origin of Species and Its Critics, 1859–1882. Philadelphia: Temple University Press, 1970.

Young, Robert M. Darwin’s Metaphor: Nature’s Place in Victorian Culture. Cambridge, U.K.: Cambridge University Press, 1985.

Peter J. Bowler

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"Darwin, Charles Robert." Complete Dictionary of Scientific Biography. 2008. Encyclopedia.com. 26 May. 2016 <http://www.encyclopedia.com>.

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Darwin, Charles

Darwin, Charles 1809-1882

DARWINS FINCHES AND THE LOGIC OF NATURAL SELECTION

SOME MISCONCEPTIONS ABOUT DARWINS THEORY

EVOLUTIONARY PSYCHOLOGY

CONTROVERSIES ABOUT DARWINISM AND HUMAN BEHAVIOR

BIBLIOGRAPHY

Charles Robert Darwin is regarded as one of the greatest scientists who ever lived. He was the son of a wealthy provincial British doctor and entered into the study of medicine at Edinburgh University, but did not like it. He was then expected by his family to become a clergyman and graduated from Cambridge University. However, he was unenthused about pursuing a career in the church.

Fortunately for him, as well as the world of the biological sciences, he was offered an unpaid position assisting the captain of the British survey ship Beagle. From this little ship, Darwin spent the next five years exploring the natural world. His observations fostered a deep interest in biology and geology and motivated him to pursue the sciences as a career and eventually develop the theory of natural selection.

Charles Darwins theory of evolution by natural selection is the only scientific explanation for the nature and variety of life on our planet. Moreover, if life in other parts of our galaxy is governed by the same natural laws that govern it here, then Darwins logic also explains the verities of life in distant parts of our galaxy. However, it does not explain the origin of life itself. Darwins theory leaves the very beginning of lifethe formation of the first primitive organic moleculesto chemists, physicists, and maybe even theologians. This entry focuses on the logic of how natural selection shapes the tools that enable organisms to survive the challenges of their environment, on some misconceptions about the theory of evolution, and on how evolutionary theory can help us understand human behavior. Finally, it briefly mentions some of the scientific and political controversies associated with current evolutionary thinking. Those interested in the vast evidence for Darwins theory should consult Mark Pagels Encyclopedia of Evolution (2002).

DARWINS FINCHES AND THE LOGIC OF NATURAL SELECTION

The Galapagos Archipelago is a group of islands off the coast of Peru. Eons ago, a single species of finch landed on one of the islands. Because resources for survival and reproduction were limited, the finches began competing for them. Some finches migrated to other islands. Since the environments of the islands differed, various biological tools, such as the shape of the birds beaks, differed in their effectiveness for dealing with these environments. Over time, fourteen species of finches evolved in the archipelago. Although the species vary in many ways, such as in size and coloration, differences in their beaks are particularly salient. The study of how the beaks of Galapagos finches contribute to survival and reproduction in the environments of the different islands has contributed greatly to the development of evolutionary theory (Weiner 1994).

Biological tools that contribute to an individuals survival and reproduction are called adaptations. An adaptation is a traitan anatomical structure, physiological process, or behavior patternthat contributes more to an individuals ability to survive, grow, or reproduce in comparison with alternate traits in the population. The large, powerful beak of the Geospiza magnirostris finch, useful for cracking large seeds, and the small, delicate beak of Certhidea olivacea, useful for extracting insects from the bark of trees, provide examples. Natural selection is the differential contribution of offspring to the next generation by genetically different members of a population, that is, the number of progeny of finches with genes for different beaks. The logic of natural selection can be explained in terms of assumptions and inferences that follow from the assumptions (Crawford 1998).

Assumption 1: The number of descendents of organisms in a population can grow exponentially.

Assumption 2: Resources enabling individuals in a population to exist can expand only arithmetically.

Assumption 3: The size of a population of individuals remains relatively stable across time.

Inference A: Competition for survival and reproduction ensues between individuals in a population.

Assumption 4: Individuals differ on traits that enable them to survive and reproduce.

Assumption 5: Some of the variation in these traits is genetic in origin.

Inference B: There is differential contribution of offspring to the next generation by genetically different members of a population (natural selection, by definition).

Inference C: Over many generations, anatomical structures, physiological processes or behavior patterns that contribute more to individuals ability to survive, grow and reproduce in comparison with alternate traits in the population (adaptations, by definition) are created.

In summary, some feature of the environmentfor example, the arrival of a new predator or a change in climateposes a problem for organisms in a population. A solution (e.g., longer legs, thicker fur) aids survival and reproduction. The above assumptions and inferences explain how natural selection provides the solution. Preexisting adaptations, sometimes called preadaptations, provide both stepping stones and limits to the solutions natural selection can provide (Pagel 2002).

SOME MISCONCEPTIONS ABOUT DARWINS THEORY

Assumptions 1, 2, and 3 do not require nature to be red in tooth and claw, as the poet Alfred Tennyson described it in In Memoriam (1850). A variety of subtle, and not so subtle, adaptations help individuals survive and reproduce. For example, animals in herds, such as wildebeests, may minimize the relative risk of predation by maneuvering to place other animals between themselves and predators (Hamilton 1971). Many types of camouflage coloration have evolved to protect organisms from their predators (Pagel 2002).

Although genes are involved in the development of all adaptations, assumptions 4 and 5 do not require that differences between individuals in evolved traits are genetically preprogrammed. Traits whose development is influenced by environmental factors are called facultative traits. Although male white-crowned sparrows, for example, cannot learn the song of another species, when they are nestlings they must hear an adult male of their own species sing and then themselves sing as juveniles if they are to sing a full song as adults (Konishi 1965).

Organisms do not evolve to act for the good of their species or the groups with which they live (Williams 1966). Since an organisms time and energy are limited, an organism that helped a member of its species or group would pay a reproductive cost (i.e., have fewer offspring) to do the helping. Hence, one of its competitors who did not help would leave more offspring, and its kind would spread through the group or species at the expense of the helpers. However, modern Darwinists have two methods of explaining the widespread helping behavior seen in nature. The first is the evolution of cooperation, in which both parties increase their reproductive success by helping the other (Trivers 1971). The second is through helping genetic kin (Hamilton 1964). Genetic kin, such as brothers and sisters, have copies of genes that are identical to those of common ancestors. If a brother inherited a gene from his mother that predisposes him to help his sister produce offspring, that helping gene can spread through the sisters offspring (his nieces and nephews) even though it reduces the brothers own reproductive success. The reason is that the sister has a fifty-fifty chance of inheriting the same helping gene from their mother and passing it on to her children, her brothers nieces and nephews.

Finally, natural selection does not have a goal. It is a purely mechanical process: traits that contribute more to survival and reproduction spread at the expense of traits that contribute less to survival and reproduction. Humans may evolve to be godlike creatures in the distant future. However, if this happens it will not be because natural selection had the goal of producing such beings.

EVOLUTIONARY PSYCHOLOGY

Evolutionary psychology uses the principles of natural selection to understand the origin and functioning of the cognitive and emotional adaptations that helped us deal with problems in our ancestral environment, known as the environment of evolutionary adaptedness, and how those mechanisms function now (Crawford and Anderson 1989).

Mating of close genetic relatives, for example, can be detrimental to reproduction and survival because it brings together deleterious recessive alleles, such as those causing some genetic diseases, in the offspring of such mates. Some Darwinists have argued that intimate rearing of brothers and sisters during their first few years, which reduces or eliminates adult sexual attraction between them, may reflect one mechanism humans evolved to avoid this problem (Westermarck 1891). This argument is supported by evidence from: (1) boys and girls reared in the same childrens houses in Israeli kibbutzim, who rarely find each other sexually attractive as adults (Shepher 1983); (2) the reduced success of Chinese shim pau marriages, in which a genetically unrelated baby girl is adopted into a family at birth with the expectation that she will marry a son of the family at their sexual maturity (Wolf 1995); and (3) sexual attraction between adult genetic siblings who were separated at birth (Bevc and Silverman 2000).

In the upper panel of Figure 1, the assumption is that brothers and sisters with genes enabling them to develop psychological mechanisms to avoid incest had greater lifetime reproductive success across evolutionary time than those who did not. The result is the ancestral genotype for the avoidance mental mechanism. The ancestral developmental environmentintimate rearing with genetic siblingsproduced the genetically organized ancestral cognitive and emotional mechanism(s) (the ancestral phenotype) that reduced sexual attraction between adult childhood intimates. The ancestral immediate environment refers to encounters between sexually mature, ancestral, opposite-sex individuals. Finally, natural selection favored the genes that enabled the development of the avoidance mental mechanisms.

The bottom panel represents an infinitesimal segment of evolutionary timea few years in an Israeli kibbutz or a Chinese sim pau marriage or the meeting of an adult brother and sister who were reared apart from birth. In all cases, the putative adaptation functions as it evolved to function with respect to childhood intimates. However, because it functions in novel environments, its decision processes produce consequences that do not serve its evolutionary purpose. That is, it does nothing to prevent the mating of the genetic siblings, while reducing the likelihood of the success of the kibbutz and shim pau marriages.

Finally, evolutionary psychology is concerned with: (1) the problems that our hominid and primate ancestors encountered in their daily lives; (2) the psychological adaptations that natural selection shaped to help deal with those problems; and (3) the way the resulting evolved adaptations function in current environments (Crawford and Anderson, 1989).

CONTROVERSIES ABOUT DARWINISM AND HUMAN BEHAVIOR

Because of the great explanatory potential of Darwins theory of evolution, it engenders continuing scientific, religious, political, and social controversy. For example, if biologically based race, gender, and social class differences in anatomy, physiology, or behavior exist, then evolutionary theory helps explain their origin and significance (Degler 1991; Pagel 2002). However, the most salient issuethe one that underlies most controversiesis the degree of genetic specialization of evolved cognitive and emotional behaviorproducing mechanisms. If natural selection produced primarily general-purpose psychological

mechanisms, as many social anthropologists and social activists argue, then evolutionary theory is of limited use in understanding human behavior. However, if it produced genetically highly-specialized psychological mechanisms, as most evolutionary psychologists argue (Barkow et al. 1992), then it is invaluable for understanding

Table 1
Outcomes of the debate about the role of ancestrally evolved innate genetic factors in the development of psychological mechanisms
Modified from p. 10 Crawford, C. B. (2004). Public Policy and Personal Decisions: The Evolutionary Context. Chapter One of Crawford, C. B. Salmon, C. (Eds.). Evolutionary psychology, public policy, and personal decisions.
Scientific Theories about the Degree of Innate Genetic Involvement in the Development of Specialized Psychological Mechanisms Possible States of Nature: Degree of Genetic Involvement in Development of Specialized Behavior Producing Psychological Mechanisms
Low: Few Limitations on Social options High: Many Limitations on Social Options
Small role for genetic involvement in development
  • The social science model of culture (Durkheim, 1895/1982)
  • Tabula rasamind as a blank slate
  • Classic behaviorism
  • Cultural determinism
  • Post modernism
  • Social constructions
  • Cultural anthropology
Valid Outcome: Implementation realizable
  • Interchangeability of sex roles
  • Eliminating homosexuality through education works
  • Walden Two (Skinner, 1948/1976) is a harmonious society
  • Effective laws for regulating sexual/reproductive behavior
  • Russian communism workable
  • Gene therapy of no use
  • Homosexuality considered abnormal/normal
  • State supported education
  • Socialized medicine
Invalid Outcome: implementation difficultSocial Constructionists Risks
  • Attempts to equalize sex roles fails
  • Eliminating homosexuality through education fails
  • Walden Two (Skinner, 1948/1976) produces oppressive society
  • Ineffective laws for regulating sexual/reproductive behaviour
  • Russian communism fails
  • Gene therapy not tried
  • Homosexuality considered abnormal/normal
  • State supported education
  • Socialized medicine
Large role for genetic involvement in development Invalid Outcome: Implementation difficultEvolutionary Psychologists Risks Valid Outcome: Implementation realizable
  • The evolutionary approach to culture (Barkow et al., 1992)
  • Classic ethology
  • Behavior genetics
  • Evolutionary psychology
  • Sociobiology
  • Human behavioral ecology
  • Evolutionary anthropology
  • Inappropriate special schools for males/females/classes/races
  • Ineffective laws for regulating sexual/reproductive behavior
  • Communism not tried
  • American dream not tried
  • Gene/drug/physical therapy not tried
  • Homosexual marriage accepted/rejected
  • State supported education
  • Social class exploitation
  • Socialized medicine
  • Appropriate special schools for males/females/classes/races
  • Effective laws for regulating sexual/reproductive behavior
  • Communism leads to corruption
  • American dream for all fails
  • Gene/drug/physical psychotherapy useful
  • Homosexual marriage accepted/rejected
  • State supported education
  • Social class exploitation
  • Socialized medicine

human mind and behavior and in developing solutions to social problems.

Table 1 shows the consequences for social policy of these two perspectives. The assumption underlying the table is that although specialized peripheral, information-processing mechanisms produce behavior, these specialized mechanisms can be assumed to develop from either a low or a high degree of innate genetic specialization of development. The columns under Possible States of Nature describe the possible states of the ancestral genotype shown in Figure 1. The rows indicate the two approaches to developing scientific explanations. Note that several different scientific approaches are listed in each row. The four cells in the table enumerate the outcomes of pairs of possibilities, that is, of pairing a high degree of genetic specialization with the scientific belief in a low degree of genetic specialization. Two cells describe valid outcomes. The two cells labeled Evolutionary Psychologists Risk and Social Constructionists Risk describe invalid outcomes. Many social activists, such as feminists and social constructionists, assume that the consequences of either making an evolutionary psychologists error or living in a society where ancestral evolved adaptations have an impact on current life, liberty, and happiness are so grave that they reject the possibility of the scientific explanations in the bottom row of the table.

Evolutionary psychologists reject this view. First, they worry about the suffering that could be caused by the social constructionist errors shown in Table 1. Second, they claim that accurate scientific knowledge about the human mind is crucial for developing more caring and harmonious societies. Third, they claim that natural selection has given the human species many evolved cognitive and emotional mechanisms, such as those underlying reciprocity and kinship, which if their functioning is understood can help us produce a better world. Fourth, they claim that evolutionary psychology is in fact an environmentalist discipline (Crawford and Anderson 1989)the specialized psychological mechanisms that produce behavior, described in Figure 1, evolved to help humans deal with problems and stresses in their environments. Hence, understanding how these evolved cognitive and emotional mechanisms work can help us create better places to pursue life, liberty, and happiness.

SEE ALSO Darwinism, Social; Natural Selection

BIBLIOGRAPHY

Barkow, Jerome, Leda Cosmides, and John Tooby. 1992. The Adapted Mind: Evolutionary Psychology and the Generation of Culture. New York: Oxford University Press.

Bevc, Irene, and Irwin Silverman. 2000. Early Separation and Sibling Incest: A Test of the Revised Westermarck Theory. Evolution and Human Behavior 21: 151-161.

Crawford, Charles. 1998. The Theory of Evolution in the Study of Human Behavior: An Introduction and Overview. In Handbook of Evolutionary Psychology: Ideas, Issues, and Applications, eds. Charles Crawford and Dennis Krebs, 3-42. Mahwah, NJ: Erlbaum.

Crawford, Charles, and Catherine Salmon. 2004. The Essence of Evolutionary Psychology: An Introduction. In Evolutionary Psychology, Public Policy, and Personal Decisions, eds. Charles Crawford and Catherine Salmon, 23-50. Mahwah, NJ: Erlbaum.

Crawford, Charles, and Judith Anderson. 1989. Sociobiology: An Environmentalist Discipline? American Psychologist 44: 1449-1459.

Degler, Carl. 1991. In Search of Human Nature: The Decline and Revival of Darwinism in American Social Thought. New York: Oxford University Press.

Hamilton, William D. 1964. The Genetical Evolution of Social Behavior, I and II. Journal of Theoretical Biology 7: 1-52.

Hamilton, William D. 1971. Geometry of the Selfish Herd. Journal of Theoretical Biology 31: 295-311.

Konishi, M. 1965. The Role of Auditory Feedback in the Control of Vocalization in the White-Crowned Sparrow. Zeitschrift für Tierpsychologie 22: 770-783.

Pagel, Mark. 2002. Encyclopedia of Evolution. Oxford: Oxford University Press.

Shepher, Joseph. 1983. Incest: A Biosocial View. New York: Academic Press.

Trivers, Robert L. 1971. The Evolution of Reciprocal Altruism. The Quarterly Review of Biology 46: 35-57.

Weiner, Jonathan. 1994. The Beak of the Finch: A Story of Evolution in our Time. New York: Knopf.

Westermarck, Edward A. 1891. The History of Human Marriage. New York: Macmillan.

Williams, George C. 1966. Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought. Princeton, NJ: Princeton University Press.

Wolf, Arthur P. 1995. Sexual Attraction and Childhood Association: A Chinese Brief for Edward Westermarck. Stanford, CA: Stanford University Press.

Charles Crawford

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Darwin, Charles

Darwin, Charles


Author of the Origin of Species (1859) and the Descent of Man (1871), Charles Darwin (18091882) famously challenged the popular belief that every species had been separately and immediately created by divine fiat. His theory of evolution by natural selection was based on what he considered an empirical fact: the presence of variation among members of every species. Darwin's powerful argument was that, in competition for limited resources, those variants having characteristics that favored them in their struggle would tend to be preserved and produce more offspring than those less advantaged. Over many generations the gradual accumulation of advantageous variations would lead to the emergence of a new species markedly different from its progenitor. Applied to humankind the argument was particularly contentious for the continuity it affirmed between animals and humans, and because the idea of species transformation was often associated with political radicalism and materialism. Darwin himself recalled that admitting the mutability of species had been like confessing to murder.

Providing a naturalistic account of species production and then of human evolution, Darwin risked offending the piety of those, including his own wife Emma Wedgwood, who wished to give the moral sense a transcendental significance. If humans had evolved from humbler species could humans be said to be made in the image of God? Was it possible to speak of an immortal soul? What remained of the argument for design, which in Christian natural theology had often presupposed the perfect adaptation of organic structures to the needs of the organism that possessed them?

Darwin was not the atheist vilified in ultraconservative religious literature, but he did become increasingly agnostic. Attacked in the name of religious orthodoxy, he found it "ludicrous" that he had once intended to become a clergyman. This was a reference to his Cambridge education, which had followed an abortive preparation in Edinburgh for a medical career. At Cambridge, the young Darwin encountered divines such as John Henslow and Adam Sedgwick who combined scientific enthusiasm with reverence for nature as a work of creation. In Edinburgh he had moved in free-thinking circles and had been introduced to the evolutionary theory of the French naturalist Jean-Baptiste Lamarck. Darwin also knew that his grandfather Erasmus Darwin had proposed organic transformation, but Charles Darwin was not yet a convert to such ideas. On leaving Cambridge his destiny would be to find ways of explaining the appearance of design in such intricate mechanisms as the human eye without recourse to the divine "Contriver" celebrated by the theologian and philosopher William Paley in his Natural Theology (1802).


Darwin's research

This destiny was shaped by a five-year voyage on which Darwin embarked in December 1831 as companion to Robert Fitzroy, captain of HMS Beagle. The ship was sailing for South America, enabling Darwin to enlarge his horizons as a naturalist and geologist. Having been captivated by the travelogues of Alexander von Humboldt he soon luxuriated in the rain forests of Brazil. As Adrian Desmond and James Moore have observed, their sublimity afforded a surrogate religious experience: "twiners entwining twiners, beautiful lepidoptera, silence, hosanna" (p. 122). Thoughts of a Christian ministry gradually receded as Darwin was enchanted by the study of nature, delighted by the discovery of fossil bones, staggered by the number of species that had become extinct. He was intrigued by resemblances between lost and living forms, by tantalizing patterns in the distribution of flora and fauna, and by disruptive natural forces. Entering the city of Concepción in Chile he found the cathedral shattered by an earthquake. At the Southern tip of South America natives of the Tierra del Fuego were struggling to survive in one of the most inhospitable regions on Earth. The world was perhaps not the "happy world" of Paley's English garden. Even before reading economist Thomas Malthus's Essay on Population in September 1838, Darwin had been "well prepared" to appreciate the struggle for existence that Malthus's arithmetic on reproductive fecundity convinced him was inexorable.

Of his visit to the Galapagos Islands it is often said that Darwin recognized that each island had its own distinctive species, eventually concluding that the different finches, for example, had diverged from a mainland ancestor, molded by nature to occupy different niches. But there was no such "Eureka" moment. Darwin had muddled his finch specimens from various islands and it was not until March 1837, following his return to England, that the ornithologist John Gould broke the exciting news that three forms of mockingbird, from different islands, were genuinely different species. Gould identified fourteen species of finch from Darwin's specimens. The enthralling question was why so many similar species lived in such proximity, but Darwin was unable to prove that the geographical isolation of each island had been responsible for the proliferation.

Darwin's earliest speculations on evolutionary change preceded his reading of Malthus. They show him playing with the idea that nature employs bisexual reproduction as a way of introducing variation into each new generation, so permitting continuing adaptation to changing conditions of existence. Darwin flirted with, but quickly rejected, the possibility of sudden mutation as a source of evolutionary change. As with the naturalist Alfred Russel Wallace later, it was when reading Malthus that the penny finally dropped and a theory of natural selection took shape.

The metaphor of "natural selection" allowed Darwin to exploit a simple analogy. Domestic animals and birds showed a degree of plasticity as breeders chose which specimens to mate when selecting for characteristics they wished to accentuate. Darwin crossed social barriers in fraternizing with pigeon fanciers and he emphasized the diversity of form ultimately derived from the common rock pigeon. Even a trained ornithologist, he argued, would be tempted to think that the pouter, runt, and fantail were not merely different varieties but different species. If, through human "selection," such effects could be produced, might not nature achieve much more in the millions of years at its disposal? For insight into the age of the Earth and for an emphasis on the incompleteness of the fossil record, which would help him to explain the absence of transitional forms in the fossil record, Darwin was indebted to the geologist Charles Lyell.


Darwin's view of religion

Did the metaphor of "selection" imply a divine selector in the management of the evolutionary process? Some of Darwin's contemporaries believed so. Darwin's own emphasis, however, was on the interplay of unconscious forces. Without denying a creator on whom the existence of everything ultimately depended, Darwin rejected the kind of deity who might be micromanaging the process. Rejecting the argument for design as formulated by Paley, Darwin's extension of natural law to explain how new species had arisen did not preclude a transcendent legislator. In his first transmutation notebook, he wrote of a "Creator who creates through laws," one who had "impressed" certain laws on nature, as a consequence of which beautiful organic forms had evolved. Darwin resembled earlier deists, admitting the existence of a creator but doubting there had been divine revelation or intervention

In certain respects his science corroded a residual faith. The more people know of the fixed laws of nature, he wrote in his Autobiography, "the more incredible do miracles become" (p. 86). As his wife recognized, the questioning mentality demanded of a scientist could induce skepticism. Debating the question whether evolution was under divine control, Darwin stressed the elements of randomness in the process. His conclusion was that the variations on which natural selection worked appeared without a prospective use in mind. The presence of so much pain and suffering also affected Darwin deeply. This was difficult to square with belief in a beneficent deity, but was consistent with his hypothesis of natural selection and with what in the first full sketch of his theory (1842) he called the "concealed war of nature."

To ascribe Darwin's agnosticism to his science would, however, be simplistic. During the Beagle voyage he was already asking himself whether an intuitive sense of God was universal among humankind, concluding it was absent among Fuegians and native Australians. Some Christian teaching he found morally repugnant. Aware of high moral standards among the freethinkers he met in the circle of the English writer and social reformer Harriet Martineau, he declared in his Autobiography that the idea of eternal damnation for those outside the fold was a "damnable doctrine" (p. 87). Although opinions differ as to when he finally renounced Christianity, the death early in 1851 of his young daughter Annie produced a crisis in which belief in a beneficent God became unsustainable. His agnosticism was to be peculiar since he retained the conviction that the universe as a whole could not be the result of chance. But so nuanced was his thinking that he came to mistrust his own conviction. If the human mind was itself the product of evolution, what guarantee was there that it could be trusted when engaging such metaphysical issues?


Religious responses to Darwin's science

Religious responses to Darwin's science have varied enormously. From 1859 until the 1930s, when a powerful new synthesis of genetics and Darwinian theory appeared, the controversial status of natural selection left plenty of scope for supplementary or alternative mechanisms for evolution in which divine control was affirmed. Strictly speaking, as the biologist Thomas Henry Huxley insisted, Darwinism had no implications for the central tenets of theism. Huxley even conceded that it was still possible to assert design in an original cosmic state from which all had developed through natural processes. Modern atheists and materialists, by contrast, frequently stress the apparently directionless aspects of biological evolution, weaving them into a completely secular and naturalistic world view.

Within the Christian churches the theory of evolution, not surprisingly, continues to be a divisive issue compounding the problems posed by historical criticism of the Bible. In some religious communities it has become the symbol of secular and liberalizing values and is still vehemently resisted. Yet religious writers have also appropriated Darwin's theory for constructive purposes, as did one of Darwin's early converts, Charles Kingsley, who concluded that a deity who could make all things make themselves exhibited greater wisdom than one who simply made things. Might a unified process of evolution testify more eloquently to a single creator than piecemeal creation? Darwin's American correspondent Asa Gray, a botanist, even suggested the theory might assist the theologians with their greatest difficultythe problem of suffering. If competition and struggle were the prerequisites of a creative process, without them there could not have been the evolutionary development that had culminated in human intelligence and responsiveness. Darwin himself had toyed with the idea that a deity who had created the possibility for humans to evolve might be considered less directly responsible for the uglier facets of nature that had also been possible in such a world. Sophisticated theologians have invoked the Darwinian theory to illuminate what they see as God's self-limitation rather than coercive agency. Others have seen in evolution evidence of divine immanence and participation in the world. It was the view of nineteenth-century Oxford theologian Aubrey Moore that, under the guise of a foe, Darwin had done the work of a friend, destroying infantile images of a conjuring god who was inactive except when intervening.

See also Creationism; Creation Science; Deism; Design; Design Argument; Divine Action; Evolution; Evolution, Biocultural; Evolution, Biological; Evolution, Human; Evolution, Theology of; Genetics; Immanence; Intelligent Design; Revelation; Scopes Trial


Bibliography

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john hedley brooke

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Charles Robert Darwin

Charles Robert Darwin

The English naturalist Charles Robert Darwin (1809-1882) discovered that natural selection was the agent for the transmutation of organisms during evolution, as did Alfred Russel Wallace independently. Darwin presented his theory in "Origin of Species."

The concept of evolution by descent dates at least from classical Greek philosophers. In the 18th century Carl Linnaeus postulated limited mutability of species by descent and hybridization. Charles Darwin's grandfather, Erasmus Darwin, and the Chevalier de Lamarck were the chief proponents of evolution about 1800. Such advocacy had little impact on the majority of naturalists, concerned to identify species, the stability of which was considered essential for their work. Natural theology regarded the perfection of adaptation between structure and mode of life in organisms as evidence for a beneficent, all-seeing, all-planning Creator. Organic structure, planned in advance for a preordained niche, was unchanged from the moment of creation. Variations in structure in these earthly imperfect versions of the Creator's idea were minor and impermanent.

In 1815 William Smith had demonstrated a sequence of fossil populations in time. Charles Lyell, adopting James Hutton's uniformitarian view that present conditions and processes were clues to the past history of the earth, wrote his Principles of Geology (1830-1833), which Darwin on his Beagle circumnavigation found most apt for his own geological observations. Fossils in South America and apparent anomalies of animal distribution triggered the task for Darwin of assembling a vast range of material. A reading of Thomas Malthus's Essay on the Principle of Population in 1838 completed Darwin's conceptual scheme.

Critics, for whom the Origin is paramount among Darwin's considerable output, have accused him of vacillation and procrastination. But recent study of unpublished manuscripts and his entire works reveal a continuity of purpose and integrity of effort to establish the high probability of the genetic relationship through descent in all forms of life. Man is dethroned as the summit of creation and as the especial concern of the Creator. This revolution in thought has had an effect on every kind of human activity.

Darwin was born on Feb. 12, 1809, at Shrewsbury, the fifth child of Robert and Susannah Darwin. His mother, who was the daughter of the famous potter Josiah Wedgwood, died when Charles was 8, and he was reared by his sisters. At the age of 9 Charles entered Shrewsbury School. His record was not outstanding, but he did learn to use English with precision and to delight in Shakespeare and Milton.

In 1825 Darwin went to Edinburgh University to study medicine. He found anatomy and materia medica dull and surgery unendurable. In 1828 he entered Christ's College, Cambridge, with the idea of taking Anglican orders. He attended John Stevens Henslow's course in botany, started a collection of beetles that became famous, and read widely. William Paley's Natural Theology (1802) delighted Darwin by its clear logical presentation, and he later regarded this study as the most worthwhile benefit from Cambridge. He received his bachelor's degree in 1831.

Voyage of the Beagle

On Henslow's recommendation Darwin was offered the position of naturalist for the second voyage of H. M. S. Beagle to survey the coast of Patagonia and Tierra del Fuego and complete observations of longitude by circumnavigation with a formidable array of chronometers. The Beagle left on Dec. 27, 1831, and returned on Oct. 2, 1836. During the voyage Darwin spent 535 days at sea and roughly 1200 on land. Enough identification of strata could be done on the spot, but sufficiently accurate identification of living organisms required systematists accessible only in London and Paris.

Darwin kept his field observations in notebooks with the specimens listed serially and their place and time of collection documented. On July 24, 1834, he wrote: "My notes are becoming bulky. I have about 600 small quarto pages full; about half of this is Geology the other imperfect descriptions of animals; with the latter I make it a rule to describe those parts which cannot be seen in specimens in spirits. I keep my private Journal distinct from the above." Toward the end of the voyage, when sea passages were long, he copied his notes and arranged them to accord with systematics, concentrating on range and habits. Geology was prepared with fewer inhibitions; he wrote from Mauritius in April 1836: "It is a rare piece of good fortune for me that of the many errant (in ships) Naturalists there have been few, or rather no, Geologists. I shall enter the field unopposed."

During the trip Darwin discovered the relevance of Lyell's uniformitarian views to the structure of St. Jago (Cape Verde Islands). He found that small locally living forms closely resembled large terrestrial fossil mammals embedded between marine shell layers and that the local sea was populated with living occupants of similar shells. He also observed the overlapping distribution on the continuous Patagonian plain of two closely related but distinct species of ostrich. An excursion along the Santa Cruz river revealed a section of strata across South America. He observed the differences between species of birds and animals on the Galápagos Islands.

Publications Resulting from Voyage

Darwin's Journal of Researches was published in 1839. With the help of a government grant toward the cost of the illustrations, the Zoology of the Voyage of the Beagle was published, in five quarto volumes, from 1839 to 1843. Specialist systematists wrote on fossil and living mammals, birds, fish, and reptiles. Darwin edited the work and contributed habits and ranges of the animals and geological notes on the fossils. Two themes run through his valuable and mostly neglected notes: distribution in space and time and observations of behavior as an aid to species diagnosis. He also published The Structure and Distribution of Coral Reefs (1842); he had studied the coral reefs in the Cocos Islands during the Beagle voyage.

Darwin abandoned the idea of fixity of species in 1837 while writing his Journal. A second edition, in 1845, had a stronger tinge of transmutation, but there was still no public avowal of the new faith. This delightful volume is his most popular and accessible work.

Darwin's Transmutation (Species) Notebooks (1837-1839) have recently been reconstructed. The notion of "selection owing to struggle" derived from his reading of Malthus in 1838. Earlier Darwin had read Pyrame de Candolle's works on plant geography, so his mind was receptive. The breadth of interest and profusion of hypotheses characteristic of Darwin, who could carry several topics in his mind at the same time, inform the whole. From this medley of facts allegedly assembled on Baconian principles all his later works derive.

It was not until Darwin's geological observations of South America were published in 1846 that he started a paper on his "first Cirripede," a shell-boring aberrant barnacle, no bigger than a pin's head, he had found at Chonos Island in 1835. This was watched while living, then dissected, and drawn while the Beagle sheltered from a week of severe storms. The working out of the relationship to other barnacles forced him to study all barnacles, a task that occupied him until 1854 and resulted in two volumes on living forms and two on fossil forms.

Darwin married Emma Wedgwood, his first cousin, in 1839. They lived in London until 1842, when ill health drove him to Down House, where he passed the rest of his life in seclusion. Four of their sons became prominent scientists: George was an astronomer and mathematician, Francis a botanist, Leonard a eugenist, and Horace a civil engineer.

Development of Ideas on Evolution

In 1842 and 1844 Darwin wrote short accounts of his transmutation views. The 1844 sketch in corrected fair copy was a testament accompanied by a letter to his wife to secure publication should he die. Late in 1844 Robert Chambers's Vestiges of Creation appeared advocating universal development by descent. A great scandal ensued, and criticism of the amateur pretensions of the author was savage. Darwin decided to bide his time and become more proficient as a biologist.

In 1855 Darwin began to study the practices of poultry and pigeon fanciers and worldwide domesticated breeds, conducted experiments on plant and animal variation and its hereditary transmission, and worried about the problem of plant and animal transport across land and water barriers, for he was persuaded of the importance of isolation for speciation. The last step in his conceptual scheme had already occurred to him in 1852 while pondering Henri Milne-Edwards's concept of diversification into specialized organs for separation of physiological functions in higher organisms and the relevance of these considerations for classification when related to the facts of embryological development. Darwin's "principle of divergence" recognizes that the dominant species must make more effective use of the territory it invades than a competing species and accordingly it becomes adapted to more diversified environments.

In May 1856 Lyell heard of Darwin's transmutation hypothesis and urged him to write an account with full references. Darwin sent the chapter on distribution to Lyell and Sir Joseph Hooker, who were deeply impressed. Darwin continued his writing, and on June 14, 1858, when he was halfway through, he received an essay from Alfred Russel Wallace containing the theory of evolution by natural selection—the same theory Darwin was working on. Lyell and Hooker arranged for a reading of a joint paper by Wallace and Darwin, and it was presented at a meeting of the Linnaean Society on July 1. The paper had little effect.

Origin of Species

On Nov. 24, 1859, Darwin published On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. The analogy of natural selection was prone to misunderstanding by readers, since it carried for them an implied purpose on the part of a "deified" Nature. Herbert Spencer's phrase "survival of the fittest" was equally misleading because the essence of Darwin's theory is that, unlike natural theology, adaptation must not be too perfect and rigid. A mutable store of variation must be available to any viable population in nature.

The publication of Darwin's book secured worldwide attention for his hypothesis and aroused impassioned controversy. His main champion was T. H. Huxley. Darwin, remote in his retreat at Down House, took painstaking note of criticism and endeavored to answer points of detail in the five more editions of Origin produced during his lifetime. He avoided trouble and made several unfortunate concessions which weakened his presentation and made his views seem vague and hesitant. The first edition is easily the best.

Later Works

In On the Various Contrivances by Which British and Foreign Orchids Are Fertilised by Insects (1862) Darwin showed how the welfare of an organism may be hidden in apparently unimportant peculiarities. It became hard to say what is "useless" in nature. His The Variation of Animals and Plants under Domestication (1868; rev. ed. 1875) expanded on a topic he had introduced in Origin. A chapter in Origin on man as the most domesticated of animals grew into the book The Descent of Man and Selection in Relation to Sex (1871). The Expression of the Emotions in Man and Animals (1872) developed from material squeezed out of the Descent.

Plants became an increasing preoccupation, the more so since Darwin had his son Francis as collaborator and amanuensis. Papers Darwin had published in 1864 were collected into The Movements and Habits of Climbing Plants (1875), and these ideas were further generalized on uniformitarian lines and published as The Power of Movement in Plants (1880). All plants, not merely climbing ones, were shown to execute to some degree exploratory "circumnutation" movements. Studies on fertilization of plants by insects recorded as early as 1840 led to The Effects of Cross and Self-Fertilisation in the Vegetable Kingdom (1876) and The Different Forms of Flowers on Plants of the Same Species (1877). Insectivorous Plants (1873) pursued the reactions of plants to stimuli. Darwin's last work returned to observations he had made in 1837: The Formation of Vegetable Mould through the Action of Worms, with Observations on Their Habits (1881). He died on April 19, 1882, and was buried in Westminster Abbey.

Further Reading

Primary sources on Darwin include The Life and Letters of Charles Darwin, edited by Francis Darwin (3 vols., 1887), which has an autobiographical chapter; More Letters of Charles Darwin, edited by Francis Darwin and A. C. Seward (1903); and his Autobiography, edited with appendix and notes by his granddaughter, Nora Barlow (1958; repr. 1969). An excellent, nontechnical account of Darwin's life and work is Sir Gavin de Beer, Charles Darwin: Evolution by Natural Selection (1964). Other biographical studies are Paul B. Sears, Charles Darwin: The Naturalist as a Cultural Force (1950), and Gerhard Wichler, Charles Darwin: The Founder of the Theory of Evolution and Natural Selection (1961). Gertrude Himmelfarb, Darwin and the Darwinian Revolution (1959), offers a provocative reinterpretation of the man and his impact.

A dramatic pictorial account of Darwin's trip around the world in the Beagle is Alan Moorehead, Darwin and the Beagle (1969), which incorporates excerpts from Darwin's autobiography, journal, and letters. Parts of Darwin's work are examined in P. R. Bell, ed., Darwin's Biological Work: Some Aspects Reconsidered (1959), and Darwin's vast influence is assessed in Michael T. Ghiselin, The Triumph of the Darwinian Method (1969). A good, succinct presentation of the essence of Darwin's ideas is Benjamin Farrington, What Darwin Really Said (1967), which can serve as a review of the major problems raised by Darwin's theories. □

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Darwin, Charles Robert (1809-1882)

Darwin, Charles Robert (1809-1882)

English naturalist

Charles Robert Darwin is credited with popularizing the concept of organic evolution by means of natural selection . Though Darwin was not the first naturalist to propose a model of biological evolution, his introduction of the mechanism of the "survival of the fittest," and discussion of the evolution of humans, marked a revolution in both science and natural philosophy.

Darwin was born in Shrewsbury, England and showed an early interest in the natural sciences, especially geology. His father, Robert Darwin, a wealthy physician, encouraged Charles to pursue studies in medicine at the University of Edinburg. Darwin soon tired of the subject, and his father sent him to Cambridge to prepare for a career in the clergy. At Cambridge, Darwin rekindled his passion for the natural sciences, often devoting more time to socializing with Cambridge scientists than to his clerical studies. With guidance from his cousin, entomologist William Darwin Fox (18051880), Darwin became increasingly involved in the growing circle of natural scientists at Cambridge. ox introduced Darwin to clergyman and biologist John Stevens Henslow (17961861). Henslow became Darwin's tutor in mathematics and theology, as well as his mentor in his personal studies of botany, geology, and zoology. Henslow profoundly influenced Darwin, and it was he who encouraged Darwin to delay seeking an appointment in the Church of England in favor of joining an expedition team and venturing overseas. After graduation, Darwin agreed to an unpaid position as naturalist aboard the H.M.S. Beagle. The expedition team was initially chartered for a three year voyage and survey of South America's Pacific coastline, but the ship pursued other ventures after their work was complete and Darwin remained part of H.M.S. Beagle' s crew for five years.

Darwin used his years aboard the Beagle to further his study of the natural sciences. In South America, Darwin became fascinated with geology. He paid close attention to changes in the land brought about by earthquakes and volcanoes. His observations led him to reject catastrophism (a theory that land forms are the result of single, catastrophic events), and instead espoused the geological theories of gradual development proposed by English geologist Charles Lyell (17971875) in his 1830 work, Principles of Geology. Yet, some of his observations in South America did not fit with Lyell's theories. Darwin disagreed with Lyell's assertion that coral reefs grew atop oceanic volcanoes and rises, and concluded that coral reefs built upon themselves. When Darwin returned to England in 1836, he and Lyell became good friends. Lyell welcomed Darwin's new research on coral reefs, and encouraged him to publish other studies from his voyages.

Darwin was elected a fellow of the Geological Society in 1836, and became a member of the Royal Society in 1839. That same year, he published his Journal of Researches into the Geology and Natural History of the Various Countries Visited by H.M.S. Beagle. Though his achievements in geology largely prompted his welcoming into Britain's scientific community, his research interests began to diverge from the discipline in the early 1840s. Discussions with other naturalists prompted Darwin's increasing interest in population diversity of fauna, extinct animals, and the presumed fixity of species. Again, he turned to notes of his observations and various specimens he gathered while on his prior expedition. The focus of his new studies was the Galápagos Islands off the Pacific coast of Ecuador. While there, Darwin was struck by the uniqueness of the island's tortoises and birds. Some neighboring islands had animal populations, which were largely similar to that of the continent, while others had seemingly different variety of species. After analyzing finch specimen from the Galápagos, Darwin concluded that species must have some means of transmutation, or ability of a species to alter over time. Darwin thus proposed that as species modified, and as old species disappeared, new varieties could be introduced. Thus, Darwin proposed an evolutionary model of animal populations.

The idea of organic evolution was not novel. French naturalist, Georges Buffon (17071788) had theorized that species were prone to development and change. Darwin's own grandfather, Erasmus Darwin, also published research regarding the evolution of species. Although the theoretical concept of evolution was not new, it remained undeveloped prior to Charles Darwin. Just as he had done with Lyell's geological theory, Darwin set about the further the understanding of evolution not merely as a philosophical concept, but as a practical scientific model for explaining the diversity of species and populations. His major contribution to the field was the introduction of a mechanism by which evolution was accomplished. Darwin believed that evolution was the product of an ongoing struggle of species to better adapt to their environment, with those that were best adapted surviving to reproduce and replace less-suited individuals. He called this phenomenon "survival of the fittest," or natural selection. In this way, Darwin believed that traits of maximum adaptiveness were transferred to future generations of the animal population, eventually resulting in new species.

Darwin finished an extensive draft of his theories in 1844, but lacked confidence in his abilities to convince others of the merits of his discoveries. Years later, prompted by rumors that a colleague was about to publish a theory similar to his own, Darwin decided to release his research. On the Origin of Species by Means of Natural Selection, or The Preservation of Favoured Races in the Struggle for Life, was published November 1859, and became an instant bestseller.

A common misconception is that On the Origin of Species was the introduction of the concept of human evolution. In fact, a discussion of human antiquity is relatively absent from the book. Darwin did not directly address the relationship between animal and human evolution until he published The Descent of Man, and Selection in Relation to Sex in 1871. Darwin introduced not only a model for the biological evolution of man, but also attempted to chart the process of man's psychological evolution. He further tried to break down the barriers between man and animals in 1872 with his work The Expression of the Emotions in Man and Animals. By observing facial features and voice sounds, Darwin asserted that man and non-human animals exhibited signs of emotion in similar ways. In the last years of his career, Darwin took the concept of organic evolution to its logical end by applying natural selection and specialization to the plant kingdom.

Darwin's works on evolution met with both debate from the scientific societies, and criticism from some members of the clergy. On the Origin of Species and The Descent of Man were both published at a time of heightened religious evangelicalism in England. Though willing to discuss his theories with colleagues in the sciences, Darwin refrained from participating in public debates concerning his research. In the last decade of his life, Darwin was disturbed about the application of his evolutionary models to social theory. By most accounts, he considered the emerging concept of the social and cultural evolution of men and civilizations, which later became known as Social Darwinism, to be a grievous misinterpretation of his works. Regardless of his opposition, he remained publicly taciturn about the impact his scientific theories on theology, scientific methodology, and social theory. Closely guarding his privacy, Darwin retired to his estate in Down. He died at Down House in 1882. Though his wishes were to receive an informal burial, Parliament immediately ordered a state burial for the famous naturalist at Westminster Abby. By the time of his death, the scientific community had largely accepted the arguments favoring his theories of evolution. Although the later discoveries in genetics and molecular biology radically reinterpreted Darwin's evolutionary mechanisms, evolutionary theory is the key and unifying theory in all biological science.

See also Evolution and evolutionary mechanisms; Evolutionary origin of bacteria and viruses

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Darwin, Charles Robert (1809-1882)

Darwin, Charles Robert (1809-1882)

English naturalist

Charles Robert Darwin is credited with popularizing the concept of organic evolution by means of natural selection. Though Darwin was not the first naturalist to propose a model of biological evolution, his introduction of the mechanism of the "survival of the fittest" and discussion of the evolution of humans, marked a revolution in both science and natural philosophy.

Darwin was born in Shrewsbury, England and showed an early interest in the natural sciences, especially geology . His father, Robert Darwin, a wealthy physician, encouraged Charles to pursue studies in medicine at the University of Edinburg. Darwin soon tired of the subject, and his father sent him to Cambridge to prepare for a career in the clergy. At Cambridge, Darwin rekindled his passion for the natural sciences, often devoting more time to socializing with Cambridge scientists than to his clerical studies. With guidance from his cousin, entomologist William Darwin Fox (18051880), Darwin became increasingly involved in the growing circle of natural scientists at Cambridge. Fox introduced Darwin to clergyman and biologist John Stevens Henslow (17961861). Henslow became Darwin's tutor in mathematics and theology, as well as his mentor in his personal studies of botany, geology, and zoology. Henslow profoundly influenced Darwin, and it was he who encouraged Darwin to delay seeking an appointment in the Church of England in favor of joining an expedition team and venturing overseas. After graduation, Darwin agreed to an unpaid position as naturalist aboard the H.M.S. Beagle. The expedition team was initially chartered for a three year voyage and survey of South America's Pacific coastline, but the ship pursued other ventures after their work was complete and Darwin remained part of H.M.S. Beagle's crew for five years.

Darwin used his years aboard The Beagle to further his study of the natural sciences. In South America , Darwin became fascinated with geology. He paid close attention to changes in the land brought about by earthquakes and volcanoes. His observations led him to reject catastrophism (a theory that land forms are the result of single, catastrophic events), and instead espoused the geological theories of gradual development proposed by English geologist Charles Lyell (17971875) in his 1830 work, Principles of Geology. Yet, some of his observations in South America did not fit with Lyell's theories. Darwin disagreed with Lyell's assertion that coral reefs grew atop oceanic volcanoes and rises, and concluded that coral reefs built upon themselves. When Darwin returned to England in 1836, he and Lyell became good

friends. Lyell welcomed Darwin's new research on coral reefs, and encouraged him to publish other studies from his voyages.

Darwin was elected a fellow of the Geological Society in 1836, and became a member of the Royal Society in 1839. That same year, he published his Journal of Researches into the Geology and Natural History of the Various Countries Visited by H.M.S. Beagle. Though his achievements in geology largely prompted his welcoming into Britain's scientific community, his research interests began to diverge from the discipline in the early 1840s. Discussions with other naturalists prompted Darwin's increasing interest in population diversity of fauna, extinct animals, and the presumed static nature of species. Again, he turned to notes of his observations and various specimens he gathered while on his prior expedition. The focus of his new studies was the Galápagos Islands off the Pacific coast of Ecuador. While there, Darwin was struck by the uniqueness of the island's tortoises and birds. Some neighboring islands had animal populations that were largely similar to that of the continent, while others had seemingly different variety of species. After analyzing finch specimens from the Galapagos, Darwin concluded that species must have some means of transmutation, or ability of a species to alter over time. Darwin thus proposed that as species modified, and as old species disappeared, new varieties could be introduced. Thus, Darwin proposed an evolutionary model of animal populations.

The idea of organic evolution was not novel. French naturalist, Georges Buffon (17071788) had theorized that species were prone to development and change. Darwin's own grandfather, Erasmus Darwin, also published research regarding the evolution of species. Although the theoretical concept of evolution was not new, it remained undeveloped prior to Charles Darwin. Just as he had done with Lyell's geological theory, Darwin set about to further the understanding of evolution not merely as a philosophical concept, but as a practical scientific model for explaining the diversity of species and populations. His major contribution to the field was the introduction of a mechanism by which evolution was accomplished. Darwin believed that evolution was the product of an ongoing struggle of species to better adapt to their environment, with those that were best adapted surviving to reproduce and replace less-suited individuals. He called this phenomenon "survival of the fittest," or natural selection. In this way, Darwin believed that traits of maximum adaptability were transferred to future generations of the animal population, eventually resulting in new species.

Darwin finished an extensive draft of his theories in 1844, but lacked confidence in his abilities to convince others of the merits of his discoveries. Years later, prompted by rumors that a colleague was about to publish a theory similar to his own, Darwin decided to release his research. On the Origin of Species by Means of Natural Selection, or The Preservation of Favoured Races in the Struggle for Life was published November 1859, and became an instant bestseller.

A common misconception is that On the Origin of Species was the introduction of the concept of human evolution. In fact, a discussion of human antiquity is relatively absent from the book. Darwin did not directly address the relationship between animal and human evolution until he published The Descent of Man, and Selection in Relation to Sex in 1871. Darwin introduced not only a model for the biological evolution of man, but also attempted to chart the process of man's psychological evolution. He further tried to break down the barriers between man and animals in 1872, with his work The Expression of the Emotions in Man and Animals. By observing facial features and voice sounds, Darwin asserted that man and non-human animals exhibited signs of emotion in similar ways. In the last years of his career, Darwin took the concept of organic evolution to its logical end by applying natural selection and specialization to the plant kingdom.

Darwin's works on evolution met with both debate from the scientific societies, and criticism from some members of the clergy. On the Origin of Species and The Descent of Man were both published at a time of heightened religious evangelicalism in England. Though willing to discuss his theories with colleagues in the sciences, Darwin refrained from participating in public debates concerning his research. In the last decade of his life, Darwin was disturbed about the application of his evolutionary models to social theory. By most accounts, he considered the emerging concept of the social and cultural evolution of men and civilizations, which later became known as Social Darwinism, to be a grievous misinterpretation of his works. Regardless of his opposition, he remained publicly taciturn about the impact his scientific theories on theology, scientific methodology, and social theory. Closely guarding his privacy, Darwin retired to his estate in Down. He died at Down House in 1882. Though his wishes were to receive an informal burial, Parliament immediately ordered a state burial for the famous naturalist at Westminster Abby. By the time of his death, the scientific community had largely accepted the arguments favoring his theories of evolution. Although the later discoveries in genetics and molecular biology radically refined and reinterpreted Darwin's evolutionary mechanisms, evolutionary theory is the key and unifying theory in all biological science.

See also Evolution, evidence of; Evolutionary mechanisms

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Darwin, Charles Robert

Charles Robert Darwin

1809-1882
British naturalist whose theory of organic evolution through natural selection revolutionized science.

Charles Robert Darwin was born in Shrewsbury, England. His father was a successful provincial physician, and his grandfather, Erasmus Darwin (1731-1802), had been a distinguished intellectual figure. Young Darwin attended the Shrewsbury School, and his early failure to achieve academic distinction continued at Edinburgh University, where he studied medicine, and at Cambridge University, where he studied theology. While at Cambridge, however, Darwin enthusiastically pursued natural history as an avocation, drawing the attention of botanist John Stevens Henslow (1796-1861) and geologist Adam Sedgwick (1785-1873). In 1831, through his connection with Henslow, Darwin joined the expedition team aboard the survey ship H.M.S. Beagle headed for the coasts of South America, the Galápagos Islands, New Zealand, and Tasmania. There is some indication that Darwin went on the voyage in order to accompany Captain FitzRoy. FitzRoy, as captain, was not to socialize with the lower status crew members on the ship, and he was worried about maintaining his mental health during the long, solitary voyage. (FitzRoy later committed suicide.) During what turned out to be a five-year voyage, Darwin, a creationist, recorded his observations. Upon his return to England, Darwin developed his theory of evolution, one of the major intellectual achievements of the nineteenth century. However, because of his creationist perspective, some of the observations made during the voyage were not useful in the development of his evolutionary ideas. In 1858, when another scientist, Alfred Russell Wallace (1823-1913) shared his observations gathered in the Malay Archipelago, Darwin hastened to publish The Origin of Species to ensure his own work would receive recognition.

Darwin's theory of evolution postulates that all species on earth change over time, and that process is

governed by the principles of natural selection. These principles hold that in the struggle for existence, some individuals, because of advantageous biological adaptation , are better able to occupy effectively a given ecological niche and therefore will produce more offspring than individuals who are less able. Realizing that his theory challenged biblically oriented views about the nature and origins of humans and animals, Darwin was extremely cautious and continued his research for another 18 years before publishing it in 1859 as On the Origin of Species by Means of Natural Selection; or, the Preservation of Favoured Races in the Struggle for Life. Every copy of the book was sold on the first day of publication. Within a few years, scientists were convinced of the soundness of the theory, although popular debate about its ideological and theological implications has continued to the present.

Although psychology was one of the fields for which Darwin's theory had revolutionary implications, it was largely left to othersnotably Darwin's cousin Francis Galton to expand them publicly. However, toward the end of his career, Darwin published three books in which he explored how human mental qualities could be understood as the result of evolution. In The Descent of Man (1871), he supported the controversial position that human beings are descended from animal ancestors. In line with this idea, he argued that the mental activities of humans and animals are fundamentally similar. He identified the presence in animals of "human" qualities such as courage and devotion, and "human" emotions, including pride, jealousy , and shame. After examining these and other common mental functions, such as memory , attention , and dreaming, Darwin concluded that the mental difference between humans and the higher animals is one of degree rather than kind.

In The Expression of the Emotionsin Man and Animals (1872), Darwin posited that human emotional expressions have evolved over time because of their link with reactions that have had adaptive or survival value. For example, an animal baring its teeth in rage is literally preparing to fight; thus its emotion gives it a physical advantage. Similarly, Darwin postulated that the "fight or flight" reaction, a heightened state of nervous arousal, was a mechanism that aided survival. He also put forth that human reactions which no longer have any clear survival value probably did in the past and that the similarity of emotional expression among all known human groups suggests a common descent from an earlier pre-human ancestor.

Darwin's final contribution to psychology was the publication in 1877 of Biographical Sketch of an Infant, based on a detailed log he had kept on the development of his eldest child, who was born in 1840. This milestone in the history of child psychology was probably the first publication of its type. One seminal idea expressed in this short work is that the individual's development parallels the development of the species to which it belongs. (Darwin had earlier made a similar observation about the development of the fetus before birth.)

Darwin's work had far-reaching influences on the theory and practice of psychology. Its emphasis on the individual's adaptation to the environment helped establish the "functional" view of the mind and of human behavior, influencing such thinkers as John Dewey and James Angell (1869-1949) in the United States, who together founded the functionalist movement at the University of Chicago. Darwin's conception of the continuity between humans and other species gave the study of animal behavior a new importance. Sigmund Freud's younger colleague, George J. Romanes (1848-1894), to whom Darwin turned over his notes on animal behavior shortly before his death, established the field of comparative psychology . Paralleling the science of comparative anatomy, this field seeks to provide insights about human beings by studying the similarities and differences between human and animal psychological functioning. In addition, Darwin's principle of natural selection led to a greater interest in variation and individual differences among members of the same species.

Darwin's other books include The Variations of Animals and Plants under Domestication (1868), Insectivorous Plants (1875), and The Power of Movement in Plants (1880). He was awarded membership in the London Geological Society in 1836 and won election to the Royal Society in 1839.

Further Reading

Clark, Ronald W. The Survival of Charles Darwin: A Biography of a Man and an Idea. New York: Random House, 1984.

Darwin, Charles. The Autobiography of Charles Darwin, 1809-1882. Edited by Nora Barlow. New York: Norton, 1969.

De Beer, Gavin. Charles Darwin: Evolution by Natural Selection. London: Doubleday, 1963.

Gruber, Howard E. Darwin on Man: A Psychological Study of Scientific Creativity. London: Wildwood House, 1974.

Ridley, Mark. The Darwin Reader. New York: Norton, 1987.

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Darwin, Charles

Charles Darwin

Born: February 12, 1809
Shrewsbury, England
Died: April 19, 1882
Kent, England

English naturalist

In The Origin of Species the English naturalist Charles Darwin outlined the theory of natural selection, or "survival of the fittest," as the explanation for the changing of living beings over time.

Early life and education

Charles Robert Darwin was born on February 12, 1809, in Shrewsbury, England, the fifth child of Robert and Susannah Darwin. His father was a successful doctor, as was his grandfather, Erasmus Darwin, who had a great influence on Charles's later theories. His mother, who was the daughter of the famous pottery maker Josiah Wedgwood (17301795), died when Charles was eight. His sisters then raised him. At the age of nine Charles entered Shrewsbury School. He was not a very good student.

In 1825 Darwin went to Edinburgh University in Scotland to study medicine, but he soon realized that he was unable to even watch an operation being performed. In 1828 he entered Christ's College, Cambridge, England, to become a minister. He soon gave up that idea also, but he continued to study. He attended John Stevens Henslow's course in botany (the study of plants), started a collection of beetles that became famous, and read widely. He received his bachelor's degree in 1831.

Voyage of the Beagle

On Henslow's recommendation Darwin was offered the position of naturalist for the second voyage of H. M. S. Beagle to survey the coast of South America. The Beagle left in December 1831 and returned in October 1836. During the voyage Darwin studied many different plants and animals and collected many specimens, concentrating on location and habits. Darwin was influenced in his Beagle studies by scientist Charles Lyell's Principles of Geology (183033), which stated that present conditions and processes were clues to the Earth's past history.

Darwin noticed on the trip that certain types of organisms existed only in certain areas and that many organisms had gone through changes that made it easier for them to survive in certain environments. For example, he studied a type of bird called a finch and realized that there were over a dozen different kinds. The size and shape of the beaks of these birds differed depending on what kind of food was available in the area each lived in.

Darwin's Journal of Researches was published in 1839. With the help of a government grant to cover the cost of the illustrations, the Zoology of the Voyage of the Beagle was published in five volumes from 1839 to 1843. A number of scientists wrote articles on fossils (the preserved remains of creatures from an earlier age), living mammals, birds, fish, and reptiles. Darwin edited the work. He contributed information on the habits and ranges of the animals and made notes on the fossils. He also published The Structure and Distribution of Coral Reefs (1842), for he had studied the coral reefs in the Cocos Islands during the Beagle voyage.

Development of ideas on evolution

In 1842 and 1844 Darwin wrote short accounts of his views on evolution (change and improvement over time). However, the publication of other related works around the same time caused great controversy (dispute) and criticism of the authors, and Darwin decided the time was not yet right for him to enter the argument. He decided to wait and do more research. Darwin studied the practices of pigeon breeders, he conducted experiments on differences in plants and animals over time, and he worried about the problem of plant and animal transport across land and water barriersfor he believed in the importance of isolation for the creation of new species.

In May 1856 Lyell heard of Darwin's ideas and urged him to write an account with full references. Darwin sent a chapter to Lyell and Sir Joseph Hooker, who were deeply impressed. In June 1858, when Darwin was halfway through his writing, he received an essay from another naturalist, Alfred Russel Wallace (18231913), containing the theory of evolution by natural selectionthe same theory Darwin was working on. Lyell and Hooker arranged for a reading of a combined paper by Wallace and Darwin, and it was presented at a meeting of the Linnaean Society in London, England, on July 1. The paper had little effect.

Origin of Species

In November 1859 Darwin published On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. His basic idea was that in the struggle to survive, some organisms adapt better than others to their surroundings, and when these survivors give birth they pass their traits on to their offspring, causing species to evolve. An English philosopher (seeker of wisdom) named Herbert Spencer created the phrase "survival of the fittest" to describe this idea.

The publication of Darwin's book brought worldwide attention to his theory and created heated dispute. Darwin was aware of all the criticism he received and tried to answer it in the additional five editions of Origin that were produced during his lifetime. In these editions he wanted to avoid trouble and wound up making several changes; this weakened his presentation and made him seem unsure of his views. The first edition is easily the best.

Later works

In On the Various Contrivances by Which British and Foreign Orchids Are Fertilised by Insects (1862), Darwin showed how the survival of an organism may be dependent on seemingly unimportant qualities. It became hard to say what is "useless" in nature. In The Variation of Animals and Plants under Domestication (1868), he expanded on a topic he had introduced in Origin. With The Descent of Man and Selection in Relation to Sex (1871) Darwin caused another uproar by suggesting that humans and apes both could be traced to a common ancestor.

Darwin became increasingly interested in plants, especially since he had his son Francis to help with the work. Papers Darwin had published in 1864 were collected into The Movements and Habits of Climbing Plants (1875), and these ideas were further explained and published as The Power of Movement in Plants (1880). Darwin's last work returned to observations he had made in 1837: The Formation of Vegetable Mould through the Action of Worms, with Observations on Their Habits (1881).

Darwin had married Emma Wedgwood, his first cousin, in 1839. Four of their sons became prominent scientists. He died on April 19, 1882, and was buried in Westminster Abbey.

For More Information

Browne, Janet. Charles Darwin: The Power of Place. New York: Knopf, 2002.

Bowlby, John. Charles Darwin: A New Life. New York: W. W. Norton, 1991.

Darwin, Francis, ed. The Life and Letters of Charles Darwin. 3 vols. New York: D. Appleton, 1887. Reprint, New York: AMS Press, 1972.

Keynes, Randall. Darwin, His Daughter, and Human Evolution. New York: Riverhead Books, 2002.

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Darwin, Charles

Darwin, Charles

Charles Robert Darwin (18091882) is widely considered the greatest naturalist of the nineteenth century. His pioneering work in the theory of evolution wrought a revolution in the study of the origins and nature of plant and animal life.

The son of Robert Darwin, a prominent English physician, Charles had an early interest in natural history, especially hunting, collecting, and geology. At his father's urging, Darwin attended medical school at Edinburgh, but found that he had little interest in medicine and returned home after two years. Wanting his son to have a respectable career, Darwin's father suggested that he should become an Anglican clergyman. Because the quiet, scholarly life of the clergyman appealed to him, Darwin agreed. He completed his degree at Cambridge in 1831. While awaiting an assignment, he was recommended for the job of naturalist on the survey ship Beagle, a voyage of nearly five years. In 1859 Darwin published The Origin of Species by Means of Natural Selection based on his discoveries during this voyage.

This seminal work contained three major discoveries. First, it presented an overwhelming amount of physical evidence of Darwin's evolutionary thesis. Naturalists had observed evolutionary change since the time of the ancient Greeks, and by the mid-1800s the idea of evolution was "in the air." But it was not until Darwin published Origin that a body of empirical evidence supported the idea of evolution. Because of Darwin's thorough and compelling work, almost no biologists today doubt the reality of evolution. Second, Darwin discovered descent from common ancestry, demonstrating that all living things are related. Tracing the lineage of any two species back far enough, one can find a common ancestor. The modern fossil record and biochemical comparisons among species verify this observation. Earlier theorists such as Jean Baptiste de Lamarck had assumed that life had originated many times and that each lineage was unique and unrelated to all others. Third, Darwin discovered and described the basic mechanism by which evolution works: natural selection. Natural selection is the differential reproductive success of some individuals in a population relative to that of others.

The Darwinian mechanism is based on differential reproductive rates. First, natural populations exhibit variation in phenotype (physical makeup) from one individual to the next, and this variation is genetically determined. For example, there is considerable variation in human height, skin color, and so on, within a population.

Second, organisms are genetically programmed to reproduce. Reproduction is a very powerful biological urge, and animals will risk or even sacrifice their lives to accomplish it. Humans feel this genetic programming in several ways, as a ticking "biological clock," as parental instinct, or as an attraction to children. As a result, natural populations have a tendency to overpopulate. Biologists define "overpopulation" in terms of limiting factors that may include food, space, mates, light, and minerals. For example, if there is enough space on an island for 1,000 deer but only enough food to sustain a population of 100, then 101 deer constitutes overpopulation. The result of overpopulation is competition among the individuals of the population for the limited resources. If there are no limited resources, there is no competition. Competition results in "survival of the fittest," an unfortunate phrase that Darwin borrowed from contemporary social theorists who are now known as "social Darwinists." In Darwinian terms, however, fitness refers only to reproductive success, not to strength, size, or (in humans) economic status.

Third, some of the variants in the population are more efficient than others in exploiting the limited resources. Success in obtaining limited resources is due largely to inherited phenotype. These individuals channel more of the limited resource through themselves and are therefore able to reproduce more successfully than individuals that compete less successfully. Thus, these selected variants pass on their genes for their genotype with greater frequency than do other variants.

Fourth, the result of this selectively favored breeding is a modification of population gene frequencies over time that may cause a change in phenotype. That is, the average state of a given character undergoing selection changes from one generation to the next. If, for example, predators feed on slower antelope, the average running speed of the population will gradually increase from generation to generation. This is observed as evolution.

And lastly, the losers, those individuals less successful at exploiting limited resources and at reproduction, may die in greater numbers (if, for example, they do not find enough food) or may find an alternative to the limited resource. Galapagos finches (Darwin's finches) with thinner beaks that were less successful at eating heavy seeds often found alternative foods such as insect larvae, which are more accessible to thinner beaks. Over time, this process results in evolutionary diversification of an ancestral species into two or more progeny species, the divergence from common ancestry recognized by Darwin.

Darwin had three great accomplishments with the publication of Origin of Species in 1859. He produced an overwhelming body of physical evidence that demonstrated the fact of evolution. Darwin also discovered descent from common ancestry and, lastly, the basic mechanism by which evolution operatesnatural selection based on differential reproductive rates of individuals in a breeding population. The implications of Darwin's discoveries have profoundly influenced almost every area of knowledge from science to religion to social theory.

See also: Extinction

Bibliography

Bowlby, John. Charles Darwin, A New Life. New York: W.W. Norton and Company, 1990.

Darwin, Charles. "The Origin of Species by Means of Natural Selection." In Mortimer J. Adler ed., Great Books of the Western World, second edition. Chicago: Encyclopaedia Britannica, 1990.

Lack, David. Darwin's Finches: An Essay on the General Biological Theory of Evolution. Gloucester, MA: Peter Smith, 1968.

Skelton, Peter, ed. Evolution: A Biological and Paleontological Approach. Harlow, England: Addison-Wesley, 1993.

ALFRED R. MARTIN

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Darwin, Charles

Darwin, Charles

English naturalist 18091882

Charles Darwin was the founder of modern evolutionary thought, and the developer, along with Alfred Russel Wallace, of the theory that natural selection is a principle driving force in evolution. Darwin is generally recognized as the single greatest thinker in the history of biology, whose contributions provided the basis for understanding the immense diversity that characterizes the natural world.

Darwin was born February 12, 1809, into a wealthy English family. A lifelong interest in natural history led him to embark, at age twenty-two, on a five-year voyage to South America aboard the HMS Beagle as the ship's naturalist. Darwin collected a wealth of specimens and made observations of both living species and fossils he encountered. Darwin was particularly struck by similarities he observed between the species found on the Galapagos Islands off the western coast of South America, and species of the mainland. He also noted differences and similarities among species found on the numerous islands of the Galapagos. The evidence suggested each species had not been independently formed by the Creator, but rather had diverged from a smaller group of common ancestors.

Upon returning home, Darwin pondered these ideas in conjunction with two other streams of thought. The first was the theory of uniformatarianism of geologist Charles Lyell. Uniformatarianism held that major geologic features, such as mountains and canyons, were formed not by rapid and short-lived catastrophic events such as floods, but rather by the slow, steady action of forces such as erosion. This mechanism suggested that Earth was much older than previously believed, a fact which Darwin saw provided the requisite time for the steady accumulation of change that would turn one species into another. The second idea was from An Essay on the Principle of Population, in which economist Thomas Malthus contrasted the potential for exponential increase in human population with the much slower increase in food supply. Malthus suggested that competition, disease, war, and famine kept the human population in check. Darwin saw that this principle provided the selective force needed to bring about change in a species.

Natural Selection

Between 1837 and 1838, Darwin developed his ideas into the principle of natural selection, which combines struggle, heritable variation, and differential reproduction. He proposed that in all species, limited resources leads to a struggle for existence, either against other members of the species, or against the environment. Members of a species differ from one another, and some of those variations influence the success of an organism's struggle. Organisms with more useful variations will leave more offspring, who inherit those variations and so are better able to cope with the environment. As this process continues over time, with successive rounds of struggle, variation, and differential reproduction, the population will become increasingly well adapted to the environment. Organisms of one species who become separated into different environments, such as the birds of the Galapagos, could develop over time into separate species through the accumulation of differences best adapted for their separate environments.

For the next decade, Darwin collected evidence to support his theory, and discussed his ideas only with a small circle of colleagues. But in 1858, he learned that naturalist Alfred Russel Wallace had developed a similar theory. Urged on by friends, Darwin agreed in 1858 to jointly submit papers with Wallace to the Journal of the Linnaean Society, and this was how the world first learned of the principle of natural selection. A year later, Darwin published On the Origin of Species, or the Preservation of Favored Races in the Struggle for Life. It is no exaggeration to call this the most important book ever published in biology. In it, Darwin provided so convincing an argument for natural selection that it became widely accepted by scientists shortly after publication. With natural selection, Darwin had provided a mechanism for evolution and an explanation for the diversity of life.

Despite the power of the arguments Darwin provided, numerous problems remained with the theory, especially with the mechanism of inheritance. Critics argued that blending of traits (for instance, tall with short to give medium height) would ultimately dilute out any variations. It was not until the particulate nature of inheritance was discovered, from the work of Gregor Mendel, that this problem was resolved.

Darwin continued to explore the ramifications of natural selection in a series of books published over the next twenty years. In 1871, he published The Descent of Man, and Selection in Relation to Sex, in which he applied the theory of natural selection to argue that humans evolved from earlier apelike creatures, and suggested sexual selection as an important adjunct to natural selection.


HUXLEY, THOMAS HENRY (18251895)

English biologist and paleontologist famous for his enthusiastic public debates in support of English naturalist Charles Darwin's theory of evolution. Huxley never finished college, but went to work as an assistant surgeon aboard a British frigate. When he returned to England four years later, he had sent home so many specimens and scientific papers that he was immediately elected to the Royal Society. He never returned to college.


On his death in 1882, in recognition of his scientific achievements, Darwin was buried in Westminster Abbey, along with Isaac Newton, Michael Faraday, and other great English scientists.

see also Adaptation; Buffon, Count (Georges-Louis Leclerc); Creationism; Evolution, Evidence for; Lamarck, Jean-Baptiste; Mendel, Gregor; Natural Selection; Sexual Selection; Species

Richard Robinson

Bibliography

Browne, Janet. Charles Darwin. Princeton, NJ: Princeton University Press, 1996.

Miller, Jonathan. Darwin for Beginners. New York: Pantheon Books, 1990.

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Darwin, Charles

Darwin, Charles

Naturalist 1809-1882

Charles Darwin was born on February 12, 1809 in Shrewsbury, England. He died at the age of seventy-three on April 19, 1882. Darwin revolutionized biology with his theory of evolution by natural selection. Although others contributed to the theory of evolution, Darwin was the first to win wide approval from the world's experts in biology.

Darwin was the son of a physician and the grandson of the physician and naturalist Erasmus Darwin. In the 1790s Erasmus Darwin proposed a theory of evolution. Darwin studied medicine at Edinburgh University and religion at Cambridge University. He received a Bachelor of Arts degree from Cambridge in 1831. There, his interest in biology was encouraged by his friendship with John Stevens Henslow, who studied plants. Henslow recommended Darwin as naturalist for a scientific expedition to South America. In 1831 Darwin began the five-year journey aboard the ship HMS Beagle. He gained valuable knowledge about the plants, animals, and natural features of the lands that he visited. In 1839 Darwin married his first cousin, Emma Wedgewood. They had ten children, seven surviving past childhood.

By 1846 Darwin had published several works on his discoveries about coral reefs and volcanic islands. For his writings and other scientific activities, he became greatly respected in the scientific community. Darwin spent years developing his theory on evolution. It is believed that he waited a long time to share it with the world because he feared the religious controversy that would result. He finally was moved to go public when he received a paper from Alfred Wallace in 1858. Wallace had the same ideas that Darwin had been pondering for 20 years. Darwin and Wallace presented their theories in a joint paper to the Linnean Society in 1858. In 1859 Darwin published his famous book On the Origin of Species by Means of Natural Selection or The Preservation of Favoured Races in the Struggle for Life. While the theory was accepted quickly in most scientific circles, religious leaders strongly opposed it. Many religious persons felt that the theory was inconsistent with the biblical book of Genesis. They feared that God had no place in Darwin's world. They claimed that in Darwin's theory, man was descended from apes and not placed in a superior position over other animals.

Darwin expanded his theory in other publications. His theories included the following: (1) most evolutionary changes were very gradual, requiring millions of years; (2) natural selection was the driving force behind evolutionary change; and (3) today's millions of species branched out from a single, original life form.

Darwin's theories spurred further research in biology. Charles Darwin is considered one of the greatest figures in the history of biology.

see also Adaptation; Biological Evolution; Natural Selection.

Denise Prendergast

Bibliography

Muir, Hazel, ed. Larousse Dictionary of Scientists. New York: Larousse Kingfisher Chambers Inc., 1994.

The Year 2000 Grolier Multimedia Encyclopedia. Danbury, CT: Grolier Interactive Inc., 1999.

Internet Resources

AboutDarwin.com. <http://www.aboutdarwin.com>.

Darwin. <www.public.coe.edu>.

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Darwin, Charles Robert

Charles Robert Darwin, 1809–82, English naturalist, b. Shrewsbury; grandson of Erasmus Darwin and of Josiah Wedgwood. He firmly established the theory of organic evolution known as Darwinism. He studied medicine at Edinburgh and for the ministry at Cambridge but lost interest in both professions during the training. His interest in natural history led to his friendship with the botanist J. S. Henslow; through him came the opportunity to make a five-year cruise (1831–36) as official naturalist aboard the Beagle. This started Darwin on a career of accumulating and assimilating data that resulted in the formulation of his concept of evolution and his explication of natural and sexual selection. He spent the remainder of his life carefully and methodically working over the information from his copious notes and from every other available source.

Independently, the naturalist A. R. Wallace had worked out a concept of evolution similar to Darwin's. Wallace sent a paper outlining his theory to Darwin in 1858, and its striking coincidences with Darwin's work led Darwin's friends to move to assure that the more cautious Darwin, who had been slow to publish, would receive credit for the independence and priority of his ideas. The next year Darwin set forth the structure of his theory and massive support for it in the superbly organized On the Origin of Species, supplemented and elaborated in his many later books, notably The Descent of Man (1871). He also formulated a theory of the origin of coral reefs.

See J. T. Costa, ed, The Annotated Origin: A Facsimile of the First Edition of On the Origin of Species (2009), and G. Chancellor and J. Van Wyhe, ed. Charles Darwin's Notebooks from the Voyage of the Beagle (2009); his autobiography (ed. by N. Barlow, 1958) and Life and Letters (ed. by F. Darwin, 1887; repr. with intro. by G. G. Simpson, 1962); The Correspondence of Charles Darwin, ed. by F. Burkhardt and S. Smith (19 vol., 1985–), and letters of Darwin and Henslow, ed. by N. Barlow (1967); biographies by G. Wichler (tr. 1961), A. Desmond and J. Moore (1991), J. Browne (2 vol., 1995–2002), and P. Johnson (2012); J. Barzun, Darwin, Marx, Wagner (rev. ed. 1958);; A. Moorehead, Darwin and the Beagle (1969, rev. ed. 1979); P. Appleman, ed., Darwin (1970, repr. 1983); D. L. Hull, Darwin and His Critics (1983); R. J. Richards, Darwin and the Emergence of Evolutionary Theories of Mind and Behavior (1989); R. Dawkins, River Out of Eden (1995); D. C. Dennett, Darwin's Dangerous Idea (1995); N. Eldredge, Reinventing Darwin (1995); S. Jones, Darwin's Ghost: The Origin of Species Updated (2000); J. Browne, Darwin's Origin of Species: A Biography (2008); A. Desmond and J. Moore, Darwin's Sacred Cause (2009, repr. 2011); R. Milner, Darwin's Universe (2009); G. Levine, Darwin the Writer (2011); R. Stott, Darwin's Ghosts: The Secret History of Evolution (2012).

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Darwin, Charles Robert

DARWIN, CHARLES ROBERT

Now recognized as a towering figure in the study of biology, Charles Darwin had an undistinguished academic career during his own lifetime. Though he barely scraped through his degree at Cambridge, Darwin was interested in natural history from early childhood. From 1831 to 1836, he served as naturalist on HMS Beagle, a small ship that circumnavigated the world, surveying to enhance the quality of navigational charts and gathering scientific specimens for the advancement of natural history. Darwin's account of the voyage of the Beagle was a literary success but contained little hint of the paradigm shift in biological thought for which Darwin soon became notorious. Darwin reflected for over twenty years after returning from his travels, and before publishing On the Origin of Species by Means of Natural Selection (1859). In this and later works, Darwin developed his theory of evolution by drawing upon his empirical observations of wildlife, fossils, and the complex relationships of localized variations in the anatomy of birds, butterflies, lizards, and other animals to their environment. Darwin's theory outraged orthodox religious beliefs in the creation (based on the myths described in Genesis that God had created the world and all that lived in it in seven days). For a time, he was reviled by a large proportion of the British establishment, but his supporters, including the eminent physician and biologist Thomas Huxley (18251895), encouraged him and scientific evidence eventually prevailed. Much further support for Darwin's theory of evolution is contained in his prolific writings after the Origin of Species. Evolution can no longer be described as a mere theory. There is such a huge body of hard scientific evidence, including much recently acquired support from molecular genetics, that evolution may be considered a fundamental fact of life.

John M. Last

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Darwin, Charles Robert

Darwin, Charles Robert (1809–82) An English naturalist who is remembered mainly for his theory of evolution, which he based largely on observations made in 1832–6 during a voyage around the world on HMS Beagle, which was engaged on a mapping survey. In 1858, in collaboration with Alfred Russel Wallace (who had reached similar conclusions), he published in the third volume of the Journal of the Linnean Society a short paper, ‘On the tendency of species to form varieties: and on the perpetuation of varieties and species by natural means of selection’, and in 1859 he published a longer account in his book, On the Origin of Species by Means of Natural Selection. In this he presented powerful evidence suggesting that change (evolution) has occurred among species, and proposing natural selection as the mechanism by which it occurs. The theory may be summarized as follows:
a. The individuals of a species show variation.

b. On average, more offspring are produced than are needed to replace their parents.

c. Populations cannot expand indefinitely and, on average, population sizes remain stable.

d. Therefore there must be competition for survival.

e. Therefore the best adapted variants (the ‘fittest’) survive. Since environmental conditions change over long periods of time, a process of natural selection occurs which favours the emergence of different variants and ultimately of new species (the ‘origin of species’). This theory is known as Darwinism. The subsequent discovery of chromosomes and genes, and the development of the science of genetics, have led to a better understanding of the ways in which variation may be caused. Modified by this modern knowledge, Darwin's theory is called ‘neo-Darwinism’.

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MICHAEL ALLABY. "Darwin, Charles Robert." A Dictionary of Plant Sciences. 1998. Encyclopedia.com. 26 May. 2016 <http://www.encyclopedia.com>.

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MICHAEL ALLABY. "Darwin, Charles Robert." A Dictionary of Plant Sciences. 1998. Retrieved May 26, 2016 from Encyclopedia.com: http://www.encyclopedia.com/doc/1O7-DarwinCharlesRobert.html

Darwin, Charles Robert

Darwin, Charles Robert (1809–82) English naturalist who is remembered mainly for his theory of evolution, which he based largely on observations made in 1832–6 during a voyage around the world on HMS Beagle, which was engaged on a mapping survey. In 1858, in collaboration with Alfred Russel Wallace (who had reached similar conclusions), he published through the Linnaean Society a short paper, ‘A theory of evolution by natural selection’ and in 1859 he published a longer account in his book, On the Origin of Species by Means of Natural Selection. In this he presented powerful evidence suggesting that change (evolution) has occurred among species, and proposed natural selection as the mechanism by which it occurs. His theory may be summarized as follows: (a) the individuals of a species show variation; (b) on average, more offspring are produced than are needed to replace their parents; (c) populations cannot expand indefinitely and, on average, population sizes remain stable; (d) therefore there must be competition for survival; (e) it is the best-adapted variants (the fittest) that survive and reproduce. Since environmental conditions change over long periods of time, a process of natural selection occurs which favours the emergence of different variants and ultimately of new species (the ‘origin of species’). This theory is known as Darwinism. The subsequent discovery of chromosomes and genes, and the development of the science of genetics, have led to a better understanding of the ways in which variation may be caused. Modified by this modern knowledge, Darwin's theory is called ‘neo-Darwinism’.

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AILSA ALLABY and MICHAEL ALLABY. "Darwin, Charles Robert." A Dictionary of Earth Sciences. 1999. Encyclopedia.com. 26 May. 2016 <http://www.encyclopedia.com>.

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AILSA ALLABY and MICHAEL ALLABY. "Darwin, Charles Robert." A Dictionary of Earth Sciences. 1999. Retrieved May 26, 2016 from Encyclopedia.com: http://www.encyclopedia.com/doc/1O13-DarwinCharlesRobert.html

Darwin, Charles Robert

Darwin, Charles Robert (1809–82) An English naturalist who is remembered mainly for his theory of evolution, which he based largely on observations made in 1832–6 during a voyage around the world on HMS Beagle, which was engaged on a mapping survey. In 1858, in collaboration with Alfred Russell Wallace (who had reached similar conclusions), he published through the Linnean Society a short paper, ‘A Theory of Evolution by Natural Selection’; and in 1859 he published a longer account in his book, On the Origin of Species by Means of Natural Selection. In this he presented powerful evidence suggesting that change (evolution) has occurred among species, and proposed natural selection as the mechanism by which it occurs. The theory may be summarized as follows:
a. The individuals of a species show variation.

b. On average, more offspring are produced than are needed to replace their parents.

c. Populations cannot expand indefinitely and, on average, population sizes remain stable.

d. Therefore there must be competition for survival.

e. Therefore the best-adapted variants (the fittest) survive.
Since environmental conditions change over long periods of time, a process of natural selection occurs which favours the emergence of different variants and ultimately of new species (the ‘origin of species’). This theory is known as Darwinism. The subsequent discovery of chromosomes and genes, and the development of the science of genetics, have led to a better understanding of the ways in which variation may be caused. Modified by this modern knowledge, Darwin's theory is called ‘neo-Darwinism’.

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MICHAEL ALLABY. "Darwin, Charles Robert." A Dictionary of Zoology. 1999. Encyclopedia.com. 26 May. 2016 <http://www.encyclopedia.com>.

MICHAEL ALLABY. "Darwin, Charles Robert." A Dictionary of Zoology. 1999. Encyclopedia.com. (May 26, 2016). http://www.encyclopedia.com/doc/1O8-DarwinCharlesRobert.html

MICHAEL ALLABY. "Darwin, Charles Robert." A Dictionary of Zoology. 1999. Retrieved May 26, 2016 from Encyclopedia.com: http://www.encyclopedia.com/doc/1O8-DarwinCharlesRobert.html

Darwin, Charles

Darwin, Charles (1809–82). Darwin made evolution scientifically respectable. Intended for medicine, he took courses at Edinburgh, but dropped out unable to bear surgery. He went on to Christ's College, Cambridge, took a pass degree, and became a clergyman, reading W. Paley with pleasure. There J. S. Henslow and Adam Sedgwick directed his enthusiasm for nature into serious science and in 1831 he was offered a place as companion to the captain on HMS Beagle surveying Cape Horn. This was the kind of opportunity that Joseph Banks had enjoyed; and on the five-year voyage round the world, Darwin became a great descriptive scientist and collector. Thinking about nature's diversity, and reading T. R. Malthus, he hit upon the idea of natural selection when he got home.

Animals and plants produced more young than could survive: those better adapted to their surroundings would be ‘selected’ by nature as the stock-breeder selected the woolliest or meatiest sheep, and their offspring would diverge, inheriting characteristics. Unlike earlier theories, his grandfather's or J. B. Lamarck's, this did not involve progressive development in accordance with a plan or law; it was open-ended. Darwin spent over 20 years collecting and marshalling evidence before publishing the Origin of Species in 1859. Meanwhile, he had married his cousin Emma Wedgwood, and settled in a remote corner of Kent; his health was poor, and he led a reclusive life, writing the standard work on barnacles. Despite furious controversy, his theory prevailed, and by the end of his life he was universally recognized.

David Knight

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JOHN CANNON. "Darwin, Charles." The Oxford Companion to British History. 2002. Encyclopedia.com. 26 May. 2016 <http://www.encyclopedia.com>.

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JOHN CANNON. "Darwin, Charles." The Oxford Companion to British History. 2002. Retrieved May 26, 2016 from Encyclopedia.com: http://www.encyclopedia.com/doc/1O110-DarwinCharles.html

Darwin, Charles Robert

Darwin, Charles Robert (1809–82) English naturalist, originator of a theory of evolution based on natural selection. In 1831 he joined a round-the-world expedition on HMS Beagle. Observations made of the flora and fauna of South America (especially the Galápagos Islands) formed the basis of his work on animal variation. The development of a similar theory by A. R. Wallace led Darwin to present his ideas to the Linnean Society in 1858, and in 1859 he published The Origin of Species, one of the world's most influential science books. Darwin argued that organisms reproduce more than is necessary to replenish their population, creating competition for survival. Opposed to the ideas of Lamarck, Darwin argued that each organism was a unique combination of genetic variations. The variations that prove helpful in the struggle to survive are passed down to the offspring of the survivors. He termed this process natural selection. Darwin did not distinguish between acquired characteristics and genetic variations. Instead, neo-Darwinism supplemented his ideas with modern research into heredity, especially mutation.

http://www.aboutdarwin.com

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"Darwin, Charles Robert." World Encyclopedia. 2005. Encyclopedia.com. 26 May. 2016 <http://www.encyclopedia.com>.

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Darwin, Charles

Darwin, Charles

English Naturalist 1809-1882

Charles Darwin was probably the most influential scientist of the nineteenth century. He is best known for his revolutionary ideas that species transmute, or evolve, by means of the primary mechanism of natural selection, ideas he set forth in his great work On the Origin of Species in 1859. This work shaped intellectual, political, and philosophical attitudes of the nineteenth century and fundamentally transformed humanity's understanding of its origins in terms of natural rather than supernatural causes. Though deemed controversial, his ideas continued to hold sway through the twentieth century. For these reasons, he is generally regarded as not only one of the great figures of the history of science, but also one of the great figures of the western intellectual tradition.

Early Life and Background

Charles Robert Darwin was born on February 12, 1809, in Shrewsbury, England, into a wealthy, distinguished family. His paternal grandfather was Erasmus Darwin, a physician who delved into many subjects, including poetry, and who made one of the early statements in support of transmutation , or the belief that species are changeable and not fixed. His father was Robert Waring Darwin, an equally successful physician, who practiced in Shrewsbury. His mother was Susannah Wedgwood, the daughter of the industrialist potter Josiah Wedgwood.

Charles Darwin's education began at home, largely through the efforts of his older sisters who took the place of his mother when she died prematurely. He was eventually sent to a local day school, and in 1818 he entered Shrewsbury School, one of the great English public schools of the nineteenth century. Though a poor student, Darwin developed a keen interest in natural history at a young age, and spent many of his leisure hours collecting insects and plants.

In 1825 Darwin enrolled at Edinburgh University with the intent of studying medicine. He was unsuccessful in his formal studies and developed a distaste for surgery. Two years later, he gave up his medical aspirations and left Edinburgh for Cambridge University, where he intended to study theology in the hopes that he could lead the respectable life of a country vicar. He failed to perform adequately in his formal studies, however. His one true interest remained in natural history, an interest that was encouraged by the professor of botany at Cambridge, John Stevens Henslow (1796-1861). Darwin became so enamored of this mentor that he came to be known as "the man who walks with Henslow." Another important influence on Darwin at this time was the professor of geology, Adam Sedgwick (1785-1873). Though Sedgwick provided the only formal training in science that Darwin was to receive, it was Henslow who provided directive influence in Darwin's life after he recommended him to the Admiralty, which was preparing a survey expedition to chart the coastline of South America. Darwin was to serve as companion to the captain, Robert FitzRoy, on the H.M.S. Beagle. On occasion he was to serve as ship's naturalist. The five-year voyage of the Beagle, beginning on December 31, 1831, and ending on October 2, 1836, was to prove the pivotal experience in shaping Darwin's subsequent scientific work. Darwin returned to England much altered from his experiences. He had not only matured during the voyage, but also distinguished himself with his superb collections, so much so that he became the center of attention in scientific circles.

Scientific Work, 1831-59

While aboard the H.M.S. Beagle, Darwin was struck by the diversity of life that he encountered and how beautifully adapted it was to diverse environments. As he traveled up and down the eastern coastline of South America he was especially interested in the geographic distribution of plants and animals that appeared to replace each other. More importantly, he was taken by the fact that extinct fossils in the area closely resembled living forms of related species, and that the flora and fauna on oceanic islands bore a striking similarity to the nearest continental landmass. He became particularly interested in the distribution of the flora and fauna of a string of recently formed volcanic islands known as the Galapagos, after the unique species of tortoises that are found there. Darwin noticed that each island in the group had an entirely unique flora and fauna. He also noticed that the species bore a striking resemblance to the species on the nearest continental landmass, the western part of South America. With insights from British geologist Sir Charles Lyell (1797-1875), who espoused a uniformitarian rather than a catastrophist geological theory, the weight of this geographic evidence suggested to Darwin that species had not been independently created. They had instead slowly diverged from preexisting species as they came to colonize new habitats and as they adapted to new ecological niches .

Although Darwin had recognized this geographic pattern, he did not have an explanation or a mechanism for how the flora and fauna were able to adapt themselves to their respective environments. The mechanism of evolution occurred to him only after his return to England and after he had been able to sort through his collections. The decisive moment came two years after his return while he was reading English economist Thomas Malthus's An Essay on the Principle of Population (1798). In this famous essay, Malthus had noticed that human populations, if left unchecked, had the tendency to double each generation. In other words, their pattern of growth was exponential. But Malthus also noted that resources necessary to sustain populations increased at a much slower rate, or arithmetically. To Malthus, this meant one thing: that at some point members of the population were subjected to strong competition for those resources, and that such events as war, disease, and famine were natural ways to keep the population in step with available resources. Darwin realized that this principle could just as easily apply to populations of animals and plants. In the competitive struggle for existence, Darwin added, those individuals who bore favorable characteristics would be more likely to survive to reproduce. They in turn were more likely to transmit these favorable characteristics to their offspring. Given enough time, subsequent generations of their offspring would depart from the parental types. Given more time, Darwin added, they would diverge even further from their ancestral types, eventually leading to new species. This principle of divergence strengthened and supported what Darwin called his theory of descent with modification by the mechanism of natural selection. It is what accounted for the origin of species.

Although he had formulated his theory between 1837 and 1838now recognized as the critical period for Darwin's intellectual development Darwin did not set these ideas formally on paper until 1842 in a rough sketch, and in 1844 in what is called his historical essay. He showed this short manuscript only to his close friend, the young botanist Joseph Dalton Hooker (1817-1911). Instead of publishing his theory, Darwin continued his research into the late 1850s, hoping to collect enough evidence to support what he knew to be a very controversial theory. Between 1837 and 1859 Darwin therefore engaged in a number of research projects intended to lend support to his theory, including some botanical studies to understand adaptation.

After formulating his theory, Darwin's plan was to collect data carefully from workers around the world to fortify his theory. In particular he was interested in both plant and animal breeding practices, which had been created by artificial selection, a form of selection through human intervention. He became especially interested in pigeon breeding, a popular hobby for Victorians, because it clearly demonstrated the stunning range of variation possible through artificial selection.

In 1839 Darwin married his first cousin, Emma Wedgwood, and in 1842 the couple left London to settle in the nearby village of Downe. There Darwin could escape from the bustle of the city and devote himself fully to his research. The quiet country environment removed from society was also important to Darwin, who began to suffer from an unknown illness, which sometimes left him incapacitated. Only a select group of scientists had access to him. By the 1850s, the inner circle of Darwin's friends, in addition to Hooker and Lyell, included a young morphologist, Thomas Henry Huxley (1825-1895), who became Darwin's intellectual defender.

Publication of On the Origin of Species

Darwin's peaceful existence was shattered in 1858 when he received a letter from a young naturalist named Alfred Russel Wallace (1823-1913). Wallace had been exploring the Malay Archipelago in Southeast Asia. Darwin faced the contents of the letter with mixed feelings: Wallace had reproduced much of Darwin's secret theory, leaving Darwin to face a possible priority dispute. At the advice of the inner circle, Darwin and Wallace published their work jointly in the Journal of the Linnaean Society in 1858. Darwin used the push from Wallace to complete a longer account of his theory, and included evidence he had gathered not only from his own observations and experiments, but also from his correspondence with naturalists and breeders from around the world. Darwin intended this work to be merely an abstract of his longer theory, but the published book was over four hundred pages. Its full title in the first 1859 edition was On the Origin of Species or the Preservation of Favored Races in the Struggle for Life. The book appeared in bookstores on November 24, 1859, and promptly sold out on the first day. It was to go through six editions.

Darwin's life was permanently changed as the theory that Huxley named "evolution" became a topic of heated debate. It was criticized on the scientific front because it failed to provide an adequate theory of heredity and because blending theories of heredity, popular at the time, would have led to a dilution of favorable characteristics. This problem was addressed only after Darwin's death by the rediscovery of Gregor Mendel and his theory of heredity in 1900. Another problem was the age of Earth, which was thought to be about four hundred million years old, an insufficient amount of time to account for the slow, gradual process that Darwin envisioned. This problem was solved after the discovery of radioactivity in the late nineteenth century that, when included in calculations, increased the age of Earth to nearly five billion years, an estimate of time long enough to account for evolution. Yet another difficulty was the fact that Darwin had no direct proof for a process that took place over a long stretch of time. Darwin knew this, and predicted that it would take some fifty years to accumulate enough evidence to support this theory. This was in fact provided beginning in 1920s with the example of industrial melanism in the peppered moth, Biston betularia.

More difficult to resolve were the theological and philosophical questions that followed from the mechanism of natural selection. Even though Darwin had only one line on human evolution in the book, the theory implied that humans were subject to the same mechanistic process as plants and animals. Mechanistic and materialistic, natural selection also challenged the argument for God's existence from design and led to a non-purposive view of the world. To some, like the poet Alfred, Lord Tennyson, a competitive nonpurposive view of nature implied that it was "red in tooth and claw."

Despite a storm of controversy over the mechanism, the fact of evolution was rapidly accepted by scientists. Only after the mechanism of heredity was understood and only after the science of genetics was integrated with natural history was the debate over the mechanism of natural selection extinguished. This did not take place until the interval of time between 1920 and 1950 and was part of the event called the evolutionary synthesis.

Botanical Work

From the start, plants figured prominently in the development of Darwin's ideas of evolution by means of natural selection. Examples from the plant world abound in On the Origin of Species and Variation of Plants and Animals Under Domestication (1868). Not only were plants easily studied and bred, but they showed a stunning assortment of adaptive features. They were to prove one of Darwin's favorite objects of study, becoming the basis of no less than seven books, most of which appeared in the latter part of Darwin's life.

Darwin's first book on plant evolution was titled Fertilization in Orchids (1862). He chose to study orchids because of the range of adaptations they displayed with respect to fertilization. Darwin recognized that these elaborate adaptations served to facilitate cross-pollination by insects such as bees. For this reason, cross-pollinated plants had flowers with bright colors and fragrant nectaries to attract bees and other insects, while wind-pollinated plants, which did not have to attract insect pollinators, had flowers with little or no color.

Darwin also observed that plants that seemed to have one or few flowers had the tendency to be hermaphrodites, having flowers of both sexes on the same plant. Bigger trees with a large number of small flowers, however, usually had flowers of only one sex. To Darwin, this implied that flowers had adapted mechanisms to ensure cross-pollination. This likely increased the variability, and also increased the vigor of the offspring. Darwin performed numerous experiments to understand the manner in which cross-pollination took place in plants and to understand the adaptive function of increased variability. In the process he noted the phenomenon of heterosis, or hybrid vigor, in the progeny of cross-pollinated plants. He also began to unravel the adaptive functions of sexual reproduction.

Darwin's work in pollination mechanisms appeared in two books. The first was Effects of Cross and Self Fertilization (1876), which was followed by Different Forms of Flowers on Plants of the Same Species (1877).

Darwin was also interested in the adaptive functions of climbing plants. He found that the phenomenon of twining , or the differential bending of plants in a clockwise or counterclockwise manner around an object, permitted young or weak plants to raise themselves higher up off the ground. This maximized exposure to air and sunlight in a relatively short time, and without the costly and time-consuming investment of woody supportive structures. The various means used by plants to climb were explored in his book Climbing Plants (1875). The mechanisms by which this and other plant movements took place was explored in Power of Movement in Plants (1880). In this book Darwin explored plant tropisms, or the manner in which plants were able to grow toward light. He determined that the stem bends toward the light because of differential growth rates: the illuminated side grew more slowly than the unilluminated side so that shoot tips appeared to bend toward the light. He postulated the existence of a substance that was diffused from the apex downward that affected growth rates. These investigations anticipated the existence and action of plant hormones.

Darwin then directed his attention to other types of movements in plants, including mechanisms for prey capture in insectivorous plants such as the sundew, Drosera rotundifolia (1880). After detailed observation and experimentation, Darwin concluded that carnivorous plants had acquired the ability to live in nitrogen-poor soil with little or no root structure by feeding on prey. In addition to developing sensory apparatus to detect and capture prey, plants had also developed a digestive system capable of breaking down proteins. Most of these observations, including experiments with the sundew plants, were the focus of his book Insectivorous Plants (1875).

Darwin's last book relating to plants was a work with a strangely ecological theme: the action of worms in turning up soil. After experiments that ran for more than fifty years, Darwin postulated that earthworms played a vital ecological role: they fed on dead leaves and other organic matter and excreted this back into the soil. In so doing, earthworms served to aerate the soil and recycle vital nutrients. These results, including quantitative estimates of how much soil was processed by earthworms, were included in The Formation of Vegetable Mould Through the Action of Worms (1881).

Darwin's botanical work is notable for its detailed observations and simple, elegant experiments. These were performed in the confines of Darwin's backyard or at greenhouses at his home in Downe. Despite the fact that some of these are now classic experiments reproduced by students the world over, they were judged harshly by the leading German plant physiologist of the late nineteenth century, Julius von Sachs (1832-1897). A revolutionary experimentalist who introduced powerful analytical laboratory methods to botanical science, launching the "New Botany," Sachs thought Darwin's naturalist tendency and simple backyard experiments to be antiquated and amateur. Nonetheless, Darwin's botanical work remains the cornerstone of his studies on variation and mechanisms of adaptation in plants and is significant for his keen insights.

The book on earthworms was published just six months before Darwin's death. Until his end, Darwin remained a productive scientist. Some of his most imaginative work was performed toward the end of his long life. His was a happy and productive life in a home filled with the voices of his ten children and numerous grandchildren. On his death in 1882, he received a rare honor for a scientist: he was given a state burial and was buried at Westminster Abbey.

see also Carnivorous Plants; Compost; Evolution of Plants; Evolution of Plants, History of; Hooker, Joseph Dalton; Mendel, Gregor; Orchidaceae; Phylogeny; Sachs, Julius von; Tropisms and Nastic Movements.

Vassiliki Betty Smocovitis

Bibliography

Allan, Mea. Darwin and His Flowers: The Key to Natural Selection. New York: Taplinger, 1977.

Bowler, Peter J. Charles Darwin: The Man and His Influence. Cambridge, UK: Cambridge University Press, 1990.

Browne, Janet. Charles Darwin: Voyaging, Vol. 1. New York: Alfred A. Knopf, 1995.

de Beer, Gavin. "Charles Darwin." In Dictionary of Scientific Biography, Vol. 3, ed. Charles Coulston Gillispie. New York: Scribner's Sons, 1970.

Desmond, Adrian, and James Moore. Darwin. London: Michael Joseph, 1991.

Huxley, Julian, and H. B. D. Kettlewell. Charles Darwin and His World. New York: Viking Press, 1965.

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Smocovitis, Vassiliki Betty. "Darwin, Charles." Plant Sciences. 2001. Encyclopedia.com. 26 May. 2016 <http://www.encyclopedia.com>.

Smocovitis, Vassiliki Betty. "Darwin, Charles." Plant Sciences. 2001. Encyclopedia.com. (May 26, 2016). http://www.encyclopedia.com/doc/1G2-3408000102.html

Smocovitis, Vassiliki Betty. "Darwin, Charles." Plant Sciences. 2001. Retrieved May 26, 2016 from Encyclopedia.com: http://www.encyclopedia.com/doc/1G2-3408000102.html

Darwin, Charles

Darwin, Charles (1809–82) British naturalist, who studied medicine in Edinburgh followed by theology at Cambridge University, intending a career in the Church. However, his interest in natural history led him to accept an invitation in 1831 to join HMS Beagle as naturalist on a round-the-world voyage. After his return five years later he published works on the geology he had observed. He was also formulating his theory of evolution by means of natural selection, but it was to be 20 years before he published The Origin of Species (1859), prompted by similar views expressed by Alfred Russel Wallace. Among his later works was The Descent of Man (1871). See also Darwinism.

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