The Variation of Animals and Plants under Domestication ... Second edition, revised, fourth thousand.

London: John Murray, 1875.

Presentation copy, inscribed in Darwin’s hand, of the second and definitive edition of the only section of Darwin’s ‘big book’ on the origin of species which was printed in his lifetime. This copy is further remarkable in having manuscript revisions, undoubtedly dictated by Darwin, in the hand of Darwin’s then amanuensis, his son Francis. These corrections were very likely for the benefit of a translator, to whom the book was presented (see below). About the first edition, published in January 1868, Francis Darwin recorded that “about half of the eight years that elapsed between its commencement and completion were spent on it. The book did not escape adverse criticism: it was said, for instance, that the public had been patiently waiting for Mr. Darwin's pièces justicatives, and that after eight years of expectation all they got was a mass of detail about pigeons, rabbits and silk worms. But the true critics welcomed it as an expansion with unrivalled wealth of illustration of a section of the Origin” (The Autobiography of Charles Darwin and Selected Letters, ed. F. Darwin, New York, 1958, p. 281). “Its two volumes were intended to provide overwhelming evidence for the ubiquity of variation … He gave numerous instances of the causes of variability, including the direct effect of the conditions of life, reversion, the effects of use and disuse, saltation, prepotency, and correlated growth. The Variation also addressed a key criticism of the Origin of Species: that it lacked an adequate understanding of inheritance” (ODNB). Along with the ascertainable facts of artificial selection, Variation also contained Darwin’s hypothesis of ‘Pangenesis,’ his hypothetical mechanism for heredity. For this second edition the text was substantially revised, with additions culled from the hundreds of letters and scores of monographs he had received over the past seven years. In this period, “Darwin eased into a more adaptationist frame of mind, suggesting that there was a role in evolutionary theory for the inheritance of acquired characteristics … Although he had never categorically excluded behaviourally or environmentally induced adaptations from his writings, he now felt that they should play a larger part” (Browne, p. 407). Most notably, Darwin modified his views on Pangenesis in ways that have been supported by recent discoveries in molecular genetics (see below). This is the final edition of the text – all subsequent editions were printed from stereotyped plates.

Provenance: Charles Darwin (1809-82) (presentation inscription on front free endpaper: ‘With very kind regards | From the Author’). There were 25 recipients of presentation copies of this second edition (Darwin Correspondence, vol. 24, p. 596f.), including the German Julius Victor Carus (1823-1903), and the Italian Giovanni Canestrini (1835-1900), the translators of the 3rd German (1878; Freeman 916) and first Italian (1876; Freeman 920) editions of Variation, respectively. It is possible that this copy is one of these: the fact that the inscription is in Charles’ hand – rather than in the hand of the publisher’s clerk as often found – suggests this is an important association copy. Moreover, the corrections correspond largely with revisions in those editions. These textual corrections are found on pp. 170, 262, 264, 425, 434 and 442 of vol. I; and in the index only, on pp. 431, 439, 450, 456 and 461, of vol. II. The hand is identifiable as that of Francis Darwin, Charles’ amanuensis at that period.

On the Origin of Species was only an abstract of the long manuscript Darwin had begun writing on 14 May 1856 which he originally intended to complete and publish as the formal presentation of his views on evolution. Compared with the Origin, this work, which was to be titled Natural Selection, has more abundant examples in illustration of Darwin's argument plus an extensive citation of sources. It had reached a length of over one quarter of a million words and was well over half completed when on 18 June 1858 Darwin’s writing was dramatically interrupted when he received an essay from Alfred Russel Wallace in Borneo entitled On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection outlining his astonishingly parallel but independently conceived theory of natural selection. Darwin felt obliged to change his plans for initial publication; and, after the brief preliminary announcement was presented jointly with Wallace’s paper at the Linnaean Society of London, he rapidly wrote out in eight months the new abstract of his views which appeared as the Origin of Species in 1859. But he still planned to publish a more extensive account of his views on evolution, and he did not abandon his long manuscript, nor write on the unused backs of the sheets for drafting other new publications as he so often did with other manuscripts.

“In the introduction [to Origin of Species, Darwin] announced that in a future publication he hoped to give ‘in detail all the facts, with references, on which my conclusions have been grounded.’ On 9 January 1860, two days after the publication of the second edition of Origin, Darwin returned to his original Natural Selection manuscript and began expanding the first two chapters on ‘Variation under Domestication.’ He had a large collection of additional notes and by the middle of June had written drafts of an introduction and two chapters on the domestication of pigeons that would eventually form part of The Variation of Animals and Plants under Domestication. Darwin apparently found writing the book tiresome and writes in his autobiography that he had been ‘tempted to publish on other subjects which at the time interested me more’ …

“Darwin continued to gather data. His own practical experiments were confined to plants but he was able to gather information from others by correspondence and even to arrange for some of his correspondents to conduct experiments on his behalf. In spite of protracted periods of illness, he made progress and in March 1865 wrote to his publisher, John Murray, saying that ‘Of present book I have 7 chapters ready for press & all others very forward, except the last & concluding one’ (the book as finally published consisted of 28 chapters). In the same letter he discussed illustrations for the book.

“Darwin had been mulling for many years on a theory of heredity. In May 1865 he sent a manuscript to his friend Thomas Huxley outlining his theory which he called pangenesis and asking whether he should publish it. In his accompanying letter Darwin wrote: ‘It is a very rash & crude hypothesis yet it has been a considerable relief to my mind, & I can hang on it a good many groups of facts.’ Huxley pointed out the similarities of pangenesis to the theories of Georges-Louis LeClerc, Comte de Buffon, and the Swiss naturalist Charles Bonnet, but eventually wrote encouraging Darwin to publish: ‘Somebody rummaging among your papers half a century hence will find Pangenesis & say ‘See this wonderful anticipation of our modern Theories—and that stupid ass, Huxley, prevented his publishing them’’.

“Just before Christmas 1866 all of the manuscript except for the final chapter was sent to the publisher. At the beginning of January, on receiving an estimate of the size of the two-volume book from the printers, he wrote to his publisher: ‘I cannot tell you how sorry I am to hear of the enormous size of my Book.’ He subsequently arranged for some of the more technical sections to be set in smaller type.

“Even at this late stage Darwin was uncertain as to whether to include a chapter on mankind. At the end of January he wrote to Murray: ‘I feel a full conviction that my Chapter on man will excite attention & plenty of abuse & I suppose abuse is as good as praise for selling a Book,’ but he then apparently decided against the idea for a week later in a letter to his close friend Joseph Hooker he explained ‘I began a chapter on Man, for which I have long collected materials, but it has grown too long, & I think I shall publish separately a very small volume, ‘An essay on the origin of mankind’’. This ‘essay’ would become two books: The Descent of Man, and Selection in Relation to Sex (1871) and The Expression of Emotions in Man and Animals (1872). The book had been advertised as early as 1865 with the unwieldy title Domesticated Animals and Cultivated Plants, or the Principles of Variation, Inheritance, Reversion, Crossing, Interbreeding, and Selection under Domestication but Darwin agreed to the shorter The Variation of Animals and Plants Under Domestication suggested by the compositors …

“Darwin received the first proofs on 1 March 1867. In the tedious task of making correction he was helped by his 23-year-old daughter Henrietta Emma Darwin. In the summer while she was away in Cornwall he wrote to commend her work, ‘All your remarks, criticisms doubts & corrections are excellent, excellent, excellent.’ While making corrections Darwin also added new material.The proofs were finished on 15 November, but there was a further delay while William Dallas prepared an index. The Variation of Animals and Plants under Domestication went on sale on 30 January 1868, thirteen years after Darwin had begun his experiments on breeding and stewing the bones of pigeons. He was feeling deflated, and concerned about how these large volumes would be received, writing: ‘if I try to read a few pages I feel fairly nauseated ... The devil take the whole book.’’ In his autobiography he estimated that he had spent 4 years 2 months ‘hard labour’ on the book.

“The first volume of The Variation of Animals and Plants under Domestication consists in a lengthy and highly detailed exploration of the mechanisms of variation, including the principle of use and disuse, the principle of the correlation of parts, and the role of the environment in causing variation, at work in a number of domestic species. Darwin starts with dogs and cats, discussing the similarities between wild and domesticated dogs, and musing on how the species changed to accommodate man's wishes. He attempts to trace a genealogy of contemporary varieties (or ‘races’) back to a few early progenitors. These arguments, as well as many others, use the vast amount of data Darwin gathered about dogs and cats to support his overarching thesis of evolution through natural selection. He then goes on to make similar points regarding horses and donkeys, sheep, goats, pigs, cattle, various types of domesticated fowl, a large number of different cultivated plants, and, most thoroughly, pigeons.

“Notably, in Chapter XXVII Darwin introduced his ‘provisional hypothesis’ of pangenesis that he had first outlined to Huxley in 1865.He proposed that each part of an organism throws off minute invisible particles which he called gemmules. These were capable of generating a similar part of an organism, thus gemmules from a foot could generate a foot. The gemmules circulated freely around the organism and could multiply by division. In sexual reproduction they were transmitted from parents to their offspring with the mixing of the gemmules producing offspring with ‘blended’ characteristics of the parents. Gemmules could also remain dormant for several generations before becoming active. He also suggested that the environment might affect the gemmules in an organism and thus allowed for the possibility of the Lamarckian inheritance of acquired characteristics.Darwin believed that his theory could explain a wide range of phenomena:

All the forms of reproduction graduate into each other and agree in their product; for it is impossible to distinguish between organisms produced from buds, from self-division, or from fertilised germs ... and as we now see that all the forms of reproduction depend on the aggregation of gemmules derived from the whole body, we can understand this general agreement. It is satisfactory to find that sexual and asexual generation ... are fundamentally the same. Parthenogenesis is no longer wonderful; in fact, the wonder is that it should not oftener occur.

“In the final pages of the book Darwin directly challenged the argument of divinely guided variation advocated by his friend and supporter the American botanist Asa Gray. He used the analogy of an architect using rocks which had broken off naturally and fallen to the foot of a cliff, asking: ‘Can it be reasonably maintained that the Creator intentionally ordered ... that certain fragments should assume certain shapes so that the builder might erect his edifice?’ In the same way, breeders or natural selection picked those that happened to be useful from variations arising by ‘general laws’, to improve plants and animals, ‘man included’. Darwin concluded with: ‘However much we may wish it, we can hardly follow Professor Asa Gray in his belief that ‘variation has been along certain beneficial lines,’ like a ‘stream along definite and useful lines of irrigation’’.Darwin confided to Hooker: ‘It is foolish to touch such subjects, but there have been so many allusions to what I think about the part which God has played in the formation of organic beings, that I thought it shabby to evade the question.’

“Darwin was concerned whether anyone would read the massive volumes and was also anxious to receive feedback from his friends on their views on pangenesis. In October 1867 before the book was published he sent copies of the corrected proofs to Asa Gray with the comment: ‘The chapter on what I call Pangenesis will be called a mad dream, and I shall be pretty well satisfied if you think it a dream worth publishing; but at the bottom of my own mind I think it contains a great truth.’ He wrote to Hooker: ‘I shall be intensely anxious to hear what you think about Pangenesis,’ and to the German naturalist Fritz Müller: ‘The greater part, as you will see, is not meant to be read; but I should very much like to hear what you think of ‘Pangenesis’.’ Few of Darwin’s colleagues shared his enthusiasm for pangenesis.Wallace was initially supportive and Darwin confided to him: ‘None of my friends will speak out, except to a certain extent Sir H. Holland, who found it very tough reading, but admits that some view ‘closely akin to it’ will have to be admitted.’

“By the end of April Variation had received more than 20 reviews.An anonymous review by George Henry Lewes in the Pall Mall Gazette praised its ‘noble calmness ... undisturbed by the heats of polemical agitation’ which made the far from calm Darwin laugh, and left him ‘cock-a-hoop’ … De Vries in 1889 praised the ‘masterly survey of the phenomena to be explained’ and accepted the idea that ‘the individual hereditary qualities of the whole organism are represented by definite material particles.’ He introduced the notion of intracellula pangenesis which, following August Weismann, rejected the idea that these particles were thrown off from all the cells of the body. He called the particles ‘pangens’, later abbreviated to ‘gene.’ In a similar vein, Weismann in his 1893 work Germ-Plasm said: ‘although Darwin modestly described his theory as a provisional hypothesis, his was, nevertheless, the first comprehensive attempt to explain all the known phenomena of heredity by a common principle ... [I]n spite of the fact that a considerable number of these assumptions are untenable, a part of the theory still remains which must be accepted as fundamental and correct − in principle at any rate − not only now but for all time to come’” (Wikipedia, accessed 16 November 2016).

In this second edition of Variation, “Darwin imagined that gemmules were ‘inconceivably minute and numerous as the stars in heaven’ and that ‘many thousand gemmules must be thrown off from the various parts of the body at each stage of development’ (p. 399). Today, we know that small RNAs [ribonucleic acids], particularly microRNAs, can be secreted from mammalian cells and circulate in blood and other body fluids. They are also capable of moving between plant cells and through the vasculature and play important roles in gene regulation, diverse cellular and developmental processes. In recent years, thousands of different RNAs have been identified in mammalian sperm, which supports Darwin's idea that ‘almost infinitely numerous and minute gemmules are contained within each bud, ovule, spermatozoon, and pollen grain’ (p. 397). Most recently, Gapp and colleagues demonstrated that stress in early life alters the production of microRNAs in the sperm of mice, which results in depressive behaviors in subsequent generations. Szyf proposed that microRNAs derived from the brains of mice that had undergone stressful experiences could make their way into the reproductive organ through the circulatory system and could then target the specific gene in sperm. Obviously, this proposal is consistent with Darwin's Pangenesis …

“Throughout his career, Darwin consistently attributed the causes of hereditary variation to changes in the environment. He clearly stated, ‘There can be no doubt that the evil effects of the long-continued exposure of the parent to injurious conditions are sometimes transmitted to the offspring’ (p. 57). In a letter to Nature, he claimed that many special fears in animals, which might be acquired through habit and the utility in, for example, predator avoidance, could be strictly inherited. His claim has now been confirmed by Dias and Ressler, who examined the inheritance of parental traumatic exposure and showed that an olfactory experience could be passed onto the progeny …

In the history of biology, neglecting certain discoveries is not uncommon. It is well known that Mendel’s experiment on plant hybridization was ignored for decades. For nearly 150 years after the formulation of Darwin’s Pangenesis, it has been resolutely excluded from the pale of biological science and is now only of historical interest. However, we can affirm that Darwin's idea that pangenetic gemmules are the molecular carriers of hereditary characters and that they are diffused through the tissues or from cell to cell has been removed from the position of a provisional hypothesis to that of a well-founded theory. It is supported by the discovery of circulating nucleic acids in human blood and plant sap and the results of experimental work in inducing hereditary changes through blood transfusion in animals and through grafting in plants. The rediscovery of Darwin's Pangenesis, if and when it happens, would, like the rediscovery of Mendel’s work, have a tremendous impact on genetics, evolution, cell biology, and the history of science” (Yongsheng Liu & Xiuju Li).

The corrections in this copy have been identified as being in the hand of Charles Darwin’s son Francis (1848-1925), who in 1874 began acting as his amanuensis. Charles was ageing and had been unwell for many years, and he now reluctantly accepted his need for a secretary, principally to respond to the enormous volume of correspondence he received. “Then Francis Darwin offered to help with the workload, ‘promising to be as civil as he could wish.’ Darwin was reluctant to relinquish that task. ‘When he did let me,’ recalled Francis, ‘he used always to say I did the civility well.’ However, in 1874, Darwin capitulated and employed Francis as his secretary and assistant … That same year Francis married Amy Ruck, the daughter of a family friend from Wales, and came to live in a house in Downe village. Francis walked up the road every day to aid his father with botanical experiments and reply to correspondents. It seems not to have occurred to Francis that Darwin was giving him employment to compensate for his failure to purse the medical profession for which he was trained” (Browne, pp. 389-390).

“The first edition in English, of 1868, was in two volumes demy octavo, the only Murray Darwin to appear in this format, and it occurs in two issues. 1,500 copies of the first were published on January 30th, having been held up for the completion of the index. Murray had sold 1,250 at his autumn sale in the previous year and Life and Letters (Vol. III, p. 99) states that the whole issue was sold out in a week … The second, of 1,250 copies, was issued in February. In the present second edition the format was reduced to the usual crown octavo. The case is in arches style, with 32 pages of inserted advertisements dated January 1876” (darwinonline). “Darwin began work on the second edition of Variation on 6 July 1875, having suggested a new edition to his publisher, John Murray, in February. However, Darwin spent much of the spring of 1875 working on Insectivorous plants, which was published in July 1875. Publication of Variation 2d ed. was initially expected in November, and then December, but was held up by floods at the printers, William Clowes & Sons. It was finally published by the second half of February 1876; although it carries an 1875 imprint, it seems that the index did not reach the printer and the number of copies to print was not decided until 1876” (Correspondence, vol. 24, Appendix III).

This copy is trimmed and in the special publisher’s presentation binding. Darwin detested having to open the top edges of his books with knives and, in his later years, demanded that his publisher produce a very small number of trimmed copies for presentation purposes. Francis Darwin wrote, “This was a favourite reform of my father’s. He wrote to the Athenaeum on the subject, Feb. 5, 1867, pointing out that a book cut, even carefully, with a paper knife collects dust on its edges far more than a machine-cut book … He tried to introduce the reform in the case of his own books but found the conservatism of booksellers too strong for him. The presentation copies of all his later books were sent out with the edges cut” (Life and Letters).

Among the other recipients of the 25 presentation copies of the second edition of Variation were Darwin’s sons Francis and George, Asa Gray, Galton, Haeckel, and Huxley.

Freeman 880; Norman 597 (for the first edition). Browne, Charles Darwin. The Power of Place, 2002. Yongsheng Liu & Xiuju Li, ‘Has Darwin's Pangenesis Been Rediscovered?’ BioScience, Vol. 64 (2014), pp. 1037–1041.



Two volumes, crown octavo (185 x 120mm), pp. xiv, 473, [1]; x, 495, [1], 32 (advertisements for John Murray's books dated January 1876), with 43 woodblocks in text (light spotting on titles). Original green cloth, arches style, with covers stamped with blind frame, gilt spines (extremities rubbed).

Item #4604

Price: $85,000.00

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