The Foundations of the Origin of Species

The writing of this Essay “during the summer of 1844,” as stated in the Autobiography30, and “from memory,” as Darwin says elsewhere31, was a remarkable achievement, and possibly renders more conceivable the still greater feat of the writing of the Origin between July 1858 and September 1859.

It is an interesting subject for speculation: what influence on the world the Essay of 1844 would have exercised, had it been published in place of the Origin. The author evidently thought of its publication in its present state as an undesirable expedient, as appears clearly from the following extracts from the Life and Letters, vol. ii. pp. 16 – 18:

C. Darwin to Mrs Darwin

Down, July 5, 1844.

“… I have just finished my sketch of my species theory. If, as I believe, my theory in time be accepted even by one competent judge, it will be a considerable step in science.

“I therefore write this in case of my sudden death, as my most solemn and last request, which I am sure you will consider the same as if legally entered in my will, that you will devote £400 to its publication, and further will yourself, or through Hensleigh32, take trouble in promoting it. I wish that my sketch be given to some competent person, with this sum to induce him to take trouble in its improvement and enlargement. I give to him all my books on Natural History, which are either scored or have references at the end to the pages, begging him carefully to look over and consider such passages as actually bearing, or by possibility bearing, on this subject. I wish you to make a list of all such books as some temptation to an editor. I also request that you will hand over «to» him all those scraps roughly divided into eight or ten brown paper portfolios. The scraps, with copied quotations from various works, are those which may aid my editor. I also request that you, or some amanuensis, will aid in deciphering any of the scraps which the editor may think possibly of use. I leave to the editor's judgment whether to interpolate these facts in the text, or as notes, or under appendices. As the looking over the references and scraps will be a long labour, and as the correcting and enlarging and altering my sketch will also take considerable time, I leave this sum of £400 as some remuneration, and any profits from the work. I consider that for this the editor is bound to get the sketch published either at a publisher's or his own risk. Many of the scraps in the portfolios contain mere rude suggestions and early views, now useless, and many of the facts will probably turn out as having no bearing on my theory.

“With respect to editors, Mr Lyell would be the best if he would undertake it; I believe he would find the work pleasant, and he would learn some facts new to him. As the editor must be a geologist as well as a naturalist, the next best editor would be Professor Forbes of London. The next best (and quite best in many respects) would be Professor Henslow. Dr Hooker would be very good. The next, Mr Strickland33. If none of these would undertake it, I would request you to consult with Mr Lyell, or some other capable man, for some editor, a geologist and naturalist. Should one other hundred pounds make the difference of procuring a good editor, I request earnestly that you will raise £500.

“My remaining collections in Natural History may be given to any one or any museum where «they» would be accepted…”

«The following note seems to have formed part of the original letter, but may have been of later date:»

“Lyell, especially with the aid of Hooker (and of any good zoological aid), would be best of all. Without an editor will pledge himself to give up time to it, it would be of no use paying such a sum.

“If there should be any difficulty in getting an editor who would go thoroughly into the subject, and think of the bearing of the passages marked in the books and copied out of scraps of paper, then let my sketch be published as it is, stating that it was done several years ago34, and from memory without consulting any works, and with no intention of publication in its present form.”

The idea that the sketch of 1844 might remain, in the event of his death, as the only record of his work, seems to have been long in his mind, for in August, 1854, when he had finished with the Cirripedes, and was thinking of beginning his “species work,” he added on the back of the above letter, “Hooker by far best man to edit my species volume. August 1854.”

I have called attention in footnotes to many points in which the Origin agrees with the Foundations. One of the most interesting is the final sentence, practically the same in the Essays of 1842 and 1844, and almost identical with the concluding words of the Origin. I have elsewhere pointed out35 that the ancestry of this eloquent passage may be traced one stage further back, – to the Note Book of 1837. I have given this sentence as an appropriate motto for the Foundations in its character of a study of general laws. It will be remembered that a corresponding motto from Whewell's Bridgewater Treatise is printed opposite the title-page of the Origin of Species.

Mr Huxley who, about the year 1887, read the Essay of 1844, remarked that “much more weight is attached to the influence of external conditions in producing variation and to the inheritance of acquired habits than in the Origin.” In the Foundations the effect of conditions is frequently mentioned, and Darwin seems to have had constantly in mind the need of referring each variation to a cause. But I gain the impression that the slighter prominence given to this view in the Origin was not due to change of opinion, but rather because he had gradually come to take this view for granted; so that in the scheme of that book, it was overshadowed by considerations which then seemed to him more pressing. With regard to the inheritance of acquired characters I am not inclined to agree with Huxley. It is certain that the Foundations contains strong recognition of the importance of germinal variation, that is of external conditions acting indirectly through the “reproductive functions.” He evidently considered this as more important than the inheritance of habit or other acquired peculiarities.

Another point of interest is the weight he attached in 1842-4 to “sports” or what are now called “mutations.” This is I think more prominent in the Foundations than in the first edition of the Origin, and certainly than in the fifth and sixth editions.

Among other interesting points may be mentioned the “good effects of crossing” being “possibly analogous to good effects of change in condition,” – a principle which he upheld on experimental grounds in his Cross and Self-Fertilisation in 1876.

In conclusion, I desire to express my thanks to Mr Wallace for a footnote he was good enough to supply: and to Professor Bateson, Sir W. Thiselton-Dyer, Dr Gadow, Professor Judd, Dr Marr, Col. Prain and Dr Stapf for information on various points. I am also indebted to Mr Rutherford, of the University Library, for his careful copy of the manuscript of 1842.

Cambridge,

June 9, 1909.

PART I

§ I. «On Variation under Domestication, and on the Principles of Selection.»

An individual organism placed under new conditions [often] sometimes varies in a small degree and in very trifling respects such as stature, fatness, sometimes colour, health, habits in animals and probably disposition. Also habits of life develope certain parts. Disuse atrophies. [Most of these slight variations tend to become hereditary.]

When the individual is multiplied for long periods by buds the variation is yet small, though greater and occasionally a single bud or individual departs widely from its type (example)36 and continues steadily to propagate, by buds, such new kind.

When the organism is bred for several generations under new or varying conditions, the variation is greater in amount and endless in kind [especially37 holds good when individuals have long been exposed to new conditions]. The nature of the external conditions tends to effect some definite change in all or greater part of offspring, – little food, small size – certain foods harmless &c. &c. organs affected and diseases – extent unknown. A certain degree of variation (Müller's twins)38 seems inevitable effect of process of reproduction. But more important is that simple «?» generation, especially under new conditions [when no crossing] «causes» infinite variation and not direct effect of external conditions, but only in as much as it affects the reproductive functions39. There seems to be no part (beau ideal of liver)40 of body, internal or external, or mind or habits, or instincts which does not vary in some small degree and [often] some «?» to a great amount.

[All such] variations [being congenital] or those very slowly acquired of all kinds [decidedly evince a tendency to become hereditary], when not so become simple variety, when it does a race. Each41 parent transmits its peculiarities, therefore if varieties allowed freely to cross, except by the chance of two characterized by same peculiarity happening to marry, such varieties will be constantly demolished42. All bisexual animals must cross, hermaphrodite plants do cross, it seems very possible that hermaphrodite animals do cross, – conclusion strengthened: ill effects of breeding in and in, good effects of crossing possibly analogous to good effects of change in condition «?»43.

Therefore if in any country or district all animals of one species be allowed freely to cross, any small tendency in them to vary will be constantly counteracted. Secondly reversion to parent form – analogue of vis medicatrix44. But if man selects, then new races rapidly formed, – of late years systematically followed, – in most ancient times often practically followed45. By such selection make race-horse, dray-horse – one cow good for tallow, another for eating &c. – one plant's good lay «illegible» in leaves another in fruit &c. &c.: the same plant to supply his wants at different times of year. By former means animals become adapted, as a direct effect to a cause, to external conditions, as size of body to amount of food. By this latter means they may also be so adapted, but further they may be adapted to ends and pursuits, which by no possibility can affect growth, as existence of tallow-chandler cannot tend to make fat. In such selected races, if not removed to new conditions, and «if» preserved from all cross, after several generations become very true, like each other and not varying. But man46 selects only «?» what is useful and curious – has bad judgment, is capricious, – grudges to destroy those that do not come up to his pattern, – has no [knowledge] power of selecting according to internal variations, – can hardly keep his conditions uniform, – [cannot] does not select those best adapted to the conditions under which «the» form «?» lives, but those most useful to him. This might all be otherwise.

§ II. «On Variation in a State of Nature and on the Natural Means of Selection.»

Let us see how far above principles of variation apply to wild animals. Wild animals vary exceedingly little – yet they are known as individuals47. British Plants, in many genera number quite uncertain of varieties and species: in shells chiefly external conditions48. Primrose and cowslip. Wild animals from different [countries can be recognized]. Specific character gives some organs as varying. Variations analogous in kind, but less in degree with domesticated animals – chiefly external and less important parts.

Our experience would lead us to expect that any and every one of these organisms would vary if «the organism were» taken away «?» and placed under new conditions. Geology proclaims a constant round of change, bringing into play, by every possible «?» change of climate and the death of pre-existing inhabitants, endless variations of new conditions. These «?» generally very slow, doubtful though «illegible» how far the slowness «?» would produce tendency to vary. But Geolog«ists» show change in configuration which, together with the accidents of air and water and the means of transportal which every being possesses, must occasionally bring, rather suddenly, organism to new conditions and «?» expose it for several generations. Hence «?» we should expect every now and then a wild form to vary49; possibly this may be cause of some species varying more than others.

According to nature of new conditions, so we might expect all or majority of organisms born under them to vary in some definite way. Further we might expect that the mould in which they are cast would likewise vary in some small degree. But is there any means of selecting those offspring which vary in the same manner, crossing them and keeping their offspring separate and thus producing selected races: otherwise as the wild animals freely cross, so must such small heterogeneous varieties be constantly counter-balanced and lost, and a uniformity of character [kept up] preserved. The former variation as the direct and necessary effects of causes, which we can see can act on them, as size of body from amount of food, effect of certain kinds of food on certain parts of bodies &c. &c.; such new varieties may then become adapted to those external [natural] agencies which act on them. But can varieties be produced adapted to end, which cannot possibly influence their structure and which it is absurd to look «at» as effects of chance. Can varieties like some vars of domesticated animals, like almost all wild species be produced adapted by exquisite means to prey on one animal or to escape from another, – or rather, as it puts out of question effects of intelligence and habits, can a plant become adapted to animals, as a plant which cannot be impregnated without agency of insect; or hooked seeds depending on animal“s existence: woolly animals cannot have any direct effect on seeds of plant. This point which all theories about climate adapting woodpecker50 to crawl «?» up trees, «illegible» miseltoe, «sentence incomplete». But if every part of a plant or animal was to vary «illegible», and if a being infinitely more sagacious than man (not an omniscient creator) during thousands and thousands of years were to select all the variations which tended towards certain ends ([or were to produce causes «?» which tended to the same end]), for instance, if he foresaw a canine animal would be better off, owing to the country producing more hares, if he were longer legged and keener sight, – greyhound produced51. If he saw that aquatic «animal would need» skinned toes. If for some unknown cause he found it would advantage a plant, which «?» like most plants is occasionally visited by bees &c.: if that plant’s seed were occasionally eaten by birds and were then carried on to rotten trees, he might select trees with fruit more agreeable to such birds as perched, to ensure their being carried to trees; if he perceived those birds more often dropped the seeds, he might well have selected a bird who would «illegible» rotten trees or [gradually select plants which «he» had proved to live on less and less rotten trees]. Who, seeing how plants vary in garden, what blind foolish man has done52 in a few years, will deny an all-seeing being in thousands of years could effect (if the Creator chose to do so), either by his own direct foresight or by intermediate means, – which will represent «?» the creator of this universe. Seems usual means. Be it remembered I have nothing to say about life and mind and all forms descending from one common type53. I speak of the variation of the existing great divisions of the organised kingdom, how far I would go, hereafter to be seen.

Before considering whether «there» be any natural means of selection, and secondly (which forms the 2nd Part of this sketch) the far more important point whether the characters and relations of animated «things» are such as favour the idea of wild species being races «?» descended from a common stock, as the varieties of potato or dahlia or cattle having so descended, let us consider probable character of [selected races] wild varieties.

Natural Selection. De Candolle’s war of nature, – seeing contented face of nature, – may be well at first doubted; we see it on borders of perpetual cold54. But considering the enormous geometrical power of increase in every organism and as «?» every country, in ordinary cases «countries» must be stocked to full extent, reflection will show that this is the case. Malthus on man, – in animals no moral [check] restraint «?» – they breed in time of year when provision most abundant, or season most favourable, every country has its seasons, – calculate robins, – oscillating from years of destruction55. If proof were wanted let any singular change of climate «occur» here «?», how astoundingly some tribes «?» increase, also introduced animals56, the pressure is always ready, – capacity of alpine plants to endure other climates, – think of endless seeds scattered abroad, – forests regaining their percentage57, – a thousand wedges58 are being forced into the œconomy of nature. This requires much reflection; study Malthus and calculate rates of increase and remember the resistance, – only periodical.

The unavoidable effect of this «is» that many of every species are destroyed either in egg or [young or mature (the former state the more common)]. In the course of a thousand generations infinitesimally small differences must inevitably tell59; when unusually cold winter, or hot or dry summer comes, then out of the whole body of individuals of any species, if there be the smallest differences in their structure, habits, instincts [senses], health &c., «it» will on an average tell; as conditions change a rather larger proportion will be preserved: so if the chief check to increase falls on seeds or eggs, so will, in the course of 1000 generations or ten thousand, those seeds (like one with down to fly60) which fly furthest and get scattered most ultimately rear most plants, and such small differences tend to be hereditary like shades of expression in human countenance. So if one parent «?» fish deposits its egg in infinitesimally different circumstances, as in rather shallower or deeper water &c., it will then «?» tell.

Let hares61 increase very slowly from change of climate affecting peculiar plants, and some other «illegible» rabbit decrease in same proportion [let this unsettle organisation of], a canine animal, who formerly derived its chief sustenance by springing on rabbits or running them by scent, must decrease too and might thus readily become exterminated. But if its form varied very slightly, the long legged fleet ones, during a thousand years being selected, and the less fleet rigidly destroyed must, if no law of nature be opposed to it, alter forms.

Remember how soon Bakewell on the same principle altered cattle and Western, sheep, – carefully avoiding a cross (pigeons) with any breed. We cannot suppose that one plant tends to vary in fruit and another in flower, and another in flower and foliage, – some have been selected for both fruit and flower: that one animal varies in its covering and another not, – another in its milk. Take any organism and ask what is it useful for and on that point it will be found to vary, – cabbages in their leaf, – corn in size «and» quality of grain, both in times of year, – kidney beans for young pod and cotton for envelope of seeds &c. &c.: dogs in intellect, courage, fleetness and smell «?»: pigeons in peculiarities approaching to monsters. This requires consideration, – should be introduced in first chapter if it holds, I believe it does. It is hypothetical at best62.

Nature’s variation far less, but such selection far more rigid and scrutinising. Man’s races not [even so well] only not better adapted to conditions than other races, but often not «?» one race adapted to its conditions, as man keeps and propagates some alpine plants in garden. Nature lets «an» animal live, till on actual proof it is found less able to do the required work to serve the desired end, man judges solely by his eye, and knows not whether nerves, muscles, arteries, are developed in proportion to the change of external form.

Besides selection by death, in bisexual animals «illegible» the selection in time of fullest vigour, namely struggle of males; even in animals which pair there seems a surplus «?» and a battle, possibly as in man more males produced than females, struggle of war or charms63. Hence that male which at that time is in fullest vigour, or best armed with arms or ornaments of its species, will gain in hundreds of generations some small advantage and transmit such characters to its offspring. So in female rearing its young, the most vigorous and skilful and industrious, «whose» instincts «are» best developed, will rear more young, probably possessing her good qualities, and a greater number will thus «be» prepared for the struggle of nature. Compared to man using a male alone of good breed. This latter section only of limited application, applies to variation of [specific] sexual characters. Introduce here contrast with Lamarck, – absurdity of habit, or chance?? or external conditions, making a woodpecker adapted to tree64.

Before considering difficulties of theory of selection let us consider character of the races produced, as now explained, by nature. Conditions have varied slowly and the organisms best adapted in their whole course of life to the changed conditions have always been selected, – man selects small dog and afterwards gives it profusion of food, – selects a long-backed and short-legged breed and gives it no particular exercise to suit this function &c. &c. In ordinary cases nature has not allowed her race to be contaminated with a cross of another race, and agriculturists know how difficult they find always to prevent this, – effect would be trueness. This character and sterility when crossed, and generally a greater amount of difference, are two main features, which distinguish domestic races from species.

[Sterility not universal admitted by all65. Gladiolus, Crinum, Calceolaria66 must be species if there be such a thing. Races of dogs and oxen: but certainly very general; indeed a gradation of sterility most perfect67 very general. Some nearest species will not cross (crocus, some heath «?»), some genera cross readily (fowls68 and grouse, peacock &c.). Hybrids no ways monstrous quite perfect except secretions69 hence even the mule has bred, – character of sterility, especially a few years ago «?» thought very much more universal than it now is, has been thought the distinguishing character; indeed it is obvious if all forms freely crossed, nature would be a chaos. But the very gradation of the character, even if it always existed in some degree which it does not, renders it impossible as marks «?» those «?» suppose distinct as species70]. Will analogy throw any light on the fact of the supposed races of nature being sterile, though none of the domestic ones are? Mr Herbert «and» Koelreuter have shown external differences will not guide one in knowing whether hybrids will be fertile or not, but the chief circumstance is constitutional differences71, such as being adapted to different climate or soil, differences which [must] probably affect the whole body of the organism and not any one part. Now wild animals, taken out of their natural conditions, seldom breed. I do not refer to shows or to Zoological Societies where many animals unite, but «do not?» breed, and others will never unite, but to wild animals caught and kept quite tame left loose and well fed about houses and living many years. Hybrids produced almost as readily as pure breds. St Hilaire great distinction of tame and domestic, – elephants, – ferrets72. Reproductive organs not subject to disease in Zoological Garden. Dissection and microscope show that hybrid is in exactly same condition as another animal in the intervals of breeding season, or those animals which taken wild and not bred in domesticity, remain without breeding their whole lives. It should be observed that so far from domesticity being unfavourable in itself «it» makes more fertile: [when animal is domesticated and breeds, productive power increased from more food and selection of fertile races]. As far as animals go might be thought «an» effect on their mind and a special case.