The Foundations of the Origin of Species

“… 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 Hensleigh[32 - Mrs Darwin’s brother.], 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 Strickland[33 - After Mr Strickland’s name comes the following sentence, which has been erased, but remains legible. “Professor Owen would be very good; but I presume he would not undertake such a work.”]. 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 ago[34 - The words “several years ago, and” seem to have been added at a later date.], 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 out[35 - Life and Letters, ii. p. 9.] 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 - Evidently a memorandum that an example should be given.] 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 [especially[37 - The importance of exposure to new conditions for several generations is insisted on in the Origin, Ed. i. p. 7, also p. 131. In the latter passage the author guards himself against the assumption that variations are “due to chance,” and speaks of “our ignorance of the cause of each particular variation.” These statements are not always remembered by his critics.] 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 - Cf. Origin, Ed. i. p. 10, vi. p. 9, “Young of the same litter, sometimes differ considerably from each other, though both the young and the parents, as Müller has remarked, have apparently been exposed to exactly the same conditions of life.”] 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 functions[39 - This is paralleled by the conclusion in the Origin, Ed. i. p. 8, that “the most frequent cause of variability may be attributed to the male and female reproductive elements having been affected prior to the act of conception.”]. There seems to be no part (beau ideal of liver)[40 - The meaning seems to be that there must be some variability in the liver otherwise anatomists would not speak of the ‘beau ideal’ of that organ.] 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. Each[41 - The position of the following passage is uncertain. “If individuals of two widely different varieties be allowed to cross, a third race will be formed – a most fertile source of the variation in domesticated animals. «In the Origin, Ed. i. p. 20 the author says that “the possibility of making distinct races by crossing has been greatly exaggerated.”» If freely allowed, the characters of pure parents will be lost, number of races thus «illegible» but differences «?» besides the «illegible». But if varieties differing in very slight respects be allowed to cross, such small variation will be destroyed, at least to our senses, – a variation [clearly] just to be distinguished by long legs will have offspring not to be so distinguished. Free crossing great agent in producing uniformity in any breed. Introduce tendency to revert to parent form.”] 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 demolished[42 - The swamping effect of intercrossing is referred to in the Origin, Ed. i. p. 103, vi. p. 126.]. 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 - A discussion on the intercrossing of hermaphrodites in relation to Knight’s views occurs in the Origin, Ed. i. p. 96, vi. p. 119. The parallelism between crossing and changed conditions is briefly given in the Origin, Ed. i. p. 267, vi. p. 391, and was finally investigated in The Effects of Cross and Self-Fertilisation in the Vegetable Kingdom, 1876.].

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 medicatrix[44 - There is an article on the vis medicatrix in Brougham’s Dissertations, 1839, a copy of which is in the author’s library.]. But if man selects, then new races rapidly formed, – of late years systematically followed, – in most ancient times often practically followed[45 - This is the classification of selection into methodical and unconscious given in the Origin, Ed. i. p. 33, vi. p. 38.]. 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 man[46 - This passage, and a similar discussion on the power of the Creator (p. 6), correspond to the comparison between the selective capacities of man and nature, in the Origin, Ed. i. p. 83, vi. p. 102.] 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 individuals[47 - i. e. they are individually distinguishable.]. British Plants, in many genera number quite uncertain of varieties and species: in shells chiefly external conditions[48 - See Origin, Ed. i. p. 133, vi. p. 165.]. 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 vary[49 - When the author wrote this sketch he seems not to have been so fully convinced of the general occurrence of variation in nature as he afterwards became. The above passage in the text possibly suggests that at this time he laid more stress on sports or mutations than was afterwards the case.]; 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 woodpecker[50 - The author may possibly have taken the case of the woodpecker from Buffon, Histoire Nat. des Oiseaux, T. vii. p. 3, 1780, where however it is treated from a different point of view. He uses it more than once, see for instance Origin, Ed. i. pp. 3, 60, 184, vi. pp. 3, 76, 220. The passage in the text corresponds with a discussion on the woodpecker and the mistletoe in Origin, Ed. i. p. 3, vi. p. 3.] 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 produced[51 - This illustration occurs in the Origin, Ed. i. pp. 90, 91, vi. pp. 110, 111.]. 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 done[52 - See Origin, Ed. i. p. 83, vi. p. 102, where the word Creator is replaced by Nature.] 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 type[53 - Note in the original. “Good place to introduce, saying reasons hereafter to be given, how far I extend theory, say to all mammalia – reasons growing weaker and weaker.”]. 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 cold[54 - See Origin, Ed. i. pp. 62, 63, vi. p. 77, where similar reference is made to De Candolle; for Malthus see Origin, p. 5.]. 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 destruction[55 - This may possibly refer to the amount of destruction going on. See Origin, Ed. i. p. 68, vi. p. 84, where there is an estimate of a later date as to death-rate of birds in winter. “Calculate robins” probably refers to a calculation of the rate of increase of birds under favourable conditions.]. If proof were wanted let any singular change of climate «occur» here «?», how astoundingly some tribes «?» increase, also introduced animals[56 - In the Origin, Ed. i. pp. 64, 65, vi. p. 80, he instances cattle and horses and certain plants in S. America and American species of plants in India, and further on, as unexpected effects of changed conditions, the enclosure of a heath, and the relation between the fertilisation of clover and the presence of cats (Origin, Ed. i. p. 74, vi. p. 91).], the pressure is always ready, – capacity of alpine plants to endure other climates, – think of endless seeds scattered abroad, – forests regaining their percentage[57 - Origin, Ed. i. p. 74, vi. p. 91. “It has been observed that the trees now growing on … ancient Indian mounds … display the same beautiful diversity and proportion of kinds as in the surrounding virgin forests.”], – a thousand wedges[58 - The simile of the wedge occurs in the Origin, Ed. i. p. 67; it is deleted in Darwin’s copy of the first edition: it does not occur in Ed. vi.] 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 tell[59 - In a rough summary at the close of the Essay, occur the words: – “Every creature lives by a struggle, smallest grain in balance must tell.”]; 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 fly[60 - Cf. Origin, Ed. i. p. 77, vi. p. 94.]) 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 hares[61 - This is a repetition of what is given at p. 6.] 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 best[62 - Compare Origin, Ed. i. p. 41, vi. p. 47. “I have seen it gravely remarked, that it was most fortunate that the strawberry began to vary just when gardeners began to attend closely to this plant. No doubt the strawberry had always varied since it was cultivated, but the slight varieties had been neglected.”].

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 charms[63 - Here we have the two types of sexual selection discussed in the Origin, Ed. i. pp. 88 et seq., vi. pp. 108 et seq.]. 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 tree[64 - It is not obvious why the author objects to “chance” or “external conditions making a woodpecker.” He allows that variation is ultimately referable to conditions and that the nature of the connexion is unknown, i.e. that the result is fortuitous. It is not clear in the original to how much of the passage the two? refer.].

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 all[65 - The meaning is “That sterility is not universal is admitted by all.”]. Gladiolus, Crinum, Calceolaria[66 - See Var. under Dom., Ed. 2, i. p. 388, where the garden forms of Gladiolus and Calceolaria are said to be derived from crosses between distinct species. Herbert’s hybrid Crinums are discussed in the Origin, Ed. i. p. 250, vi. p. 370. It is well known that the author believed in a multiple origin of domestic dogs.] must be species if there be such a thing. Races of dogs and oxen: but certainly very general; indeed a gradation of sterility most perfect[67 - The argument from gradation in sterility is given in the Origin, Ed. i. pp. 248, 255, vi. pp. 368, 375. In the Origin, I have not come across the cases mentioned, viz. crocus, heath, or grouse and fowl or peacock. For sterility between closely allied species, see Origin, Ed. i. p. 257, vi. p. 377. In the present essay the author does not distinguish between fertility between species and the fertility of the hybrid offspring, a point on which he insists in the Origin, Ed. i. p. 245, vi. p. 365.] very general. Some nearest species will not cross (crocus, some heath «?»), some genera cross readily (fowls[68 - Ackermann (Ber. d. Vereins f. Naturkunde zu Kassel, 1898, p. 23) quotes from Gloger that a cross has been effected between a domestic hen and a Tetrao tetrix; the offspring died when three days old.] and grouse, peacock &c.). Hybrids no ways monstrous quite perfect except secretions[69 - No doubt the sexual cells are meant. I do not know on what evidence it is stated that the mule has bred.] 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 species[70 - The sentence is all but illegible. I think that the author refers to forms usually ranked as varieties having been marked as species when it was found that they were sterile together. See the case of the red and blue Anagallis given from Gärtner in the Origin, Ed. i. p. 247, vi. p. 368.]]. 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 differences[71 - In the Origin, Ed. i. p. 258, where the author speaks of constitutional differences in this connexion, he specifies that they are confined to the reproductive system.], 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, – ferrets[72 - The sensitiveness of the reproductive system to changed conditions is insisted on in the Origin, Ed. i. p. 8, vi. p. 10.The ferret is mentioned, as being prolific in captivity, in Var. under Dom., Ed. 2, ii. p. 90.]. 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.

But turning to plants we find same class of facts. I do not refer to seeds not ripening, perhaps the commonest cause, but to plants not setting, which either is owing to some imperfection of ovule or pollen. Lindley says sterility is the [curse] bane of all propagators, – Linnæus about alpine plants. American bog plants, – pollen in exactly same state as in hybrids, – same in geraniums. Persian and Chinese[73 - Lindley’s remark is quoted in the Origin, Ed. i. p. 9. Linnæus’ remark is to the effect that Alpine plants tend to be sterile under cultivation (see Var. under Dom., Ed. 2, ii. p. 147). In the same place the author speaks of peat-loving plants being sterile in our gardens, – no doubt the American bog-plants referred to above. On the following page (p. 148) the sterility of the lilac (Syringa persica and chinensis) is referred to.] lilac will not seed in Italy and England. Probably double plants and all fruits owe their developed parts primarily «?» to sterility and extra food thus «?» applied[74 - The author probably means that the increase in the petals is due to a greater food supply being available for them owing to sterility. See the discussion in Var. under Dom., Ed. 2, ii. p. 151. It must be noted that doubleness of the flower may exist without noticeable sterility.]. There is here gradation «in» sterility and then parts, like diseases, are transmitted hereditarily. We cannot assign any cause why the Pontic Azalea produces plenty of pollen and not American[75 - I have not come across this case in the author’s works.], why common lilac seeds and not Persian, we see no difference in healthiness. We know not on what circumstances these facts depend, why ferret breeds, and cheetah[76 - For the somewhat doubtful case of the cheetah (Felis jubata) see Var. under Dom., Ed. 2, ii. p. 133. I do not know to what fact “pig in India” refers.], elephant and pig in India will not.

Now in crossing it is certain every peculiarity in form and constitution is transmitted: an alpine plant transmits its alpine tendency to its offspring, an American plant its American-bog constitution, and «with» animals, those peculiarities, on which[77 - This sentence should run “on which depends their incapacity to breed in unnatural conditions.”] when placed out of their natural conditions they are incapable of breeding; and moreover they transmit every part of their constitution, their respiration, their pulse, their instinct, which are all suddenly modified, can it be wondered at that they are incapable of breeding? I think it may be truly said it would be more wonderful if they did. But it may be asked why have not the recognised varieties, supposed to have been produced through the means of man, [not refused to breed] have all bred[78 - This sentence ends in confusion: it should clearly close with the words “refused to breed” in place of the bracket and the present concluding phrase.]. Variation depends on change of condition and selection[79 - The author doubtless refers to the change produced by the summation of variation by means of selection.], as far as man’s systematic or unsystematic selection «has» gone; he takes external form, has little power from ignorance over internal invisible constitutional differences. Races which have long been domesticated, and have much varied, are precisely those which were capable of bearing great changes, whose constitutions were adapted to a diversity of climates. Nature changes slowly and by degrees. According to many authors probably breeds of dogs are another case of modified species freely crossing. There is no variety which «illegible» has been «illegible» adapted to peculiar soil or situation for a thousand years and another rigorously adapted to another, till such can be produced, the question is not tried[80 - The meaning of this sentence is made clear by a passage in the MS. of 1844: – “Until man selects two varieties from the same stock, adapted to two climates or to other different external conditions, and confines each rigidly for one or several thousand years to such conditions, always selecting the individuals best adapted to them, he cannot be said to have even commenced the experiment.” That is, the attempt to produce mutually sterile domestic breeds.]. Man in past ages, could transport into different climates, animals and plants which would freely propagate in such new climates. Nature could effect, with selection, such changes slowly, so that precisely those animals which are adapted to submit to great changes have given rise to diverse races, – and indeed great doubt on this head[81 - This passage is to some extent a repetition of a previous one and may have been intended to replace an earlier sentence. I have thought it best to give both. In the Origin, Ed. i. p. 141, vi. p. 176, the author gives his opinion that the power of resisting diverse conditions, seen in man and his domestic animals, is an example “of a very common flexibility of constitution.”].

Before leaving this subject well to observe that it was shown that a certain amount of variation is consequent on mere act of reproduction, both by buds and sexually, – is vastly increased when parents exposed for some generations to new conditions[82 - In the Origin, Ed. i. Chs. I. and V., the author does not admit reproduction, apart from environment, as being a cause of variation. With regard to the cumulative effect of new conditions there are many passages in the Origin, Ed. i. e.g. pp. 7, 12, vi. pp. 8, 14.], and we now find that many animals when exposed for first time to very new conditions, are «as» incapable of breeding as hybrids. It [probably] bears also on supposed fact of crossed animals when not infertile, as in mongrels, tending to vary much, as likewise seems to be the case, when true hybrids possess just sufficient fertility to propagate with the parent breeds and inter se for some generations. This is Koelreuter’s belief. These facts throw light on each other and support the truth of each other, we see throughout a connection between the reproductive faculties and exposure to changed conditions of life whether by crossing or exposure of the individuals[83 - As already pointed out, this is the important principle investigated in the author’s Cross and Self-Fertilisation. Professor Bateson has suggested to me that the experiments should be repeated with gametically pure individuals.].

Difficulties on theory of selection[84 - In the Origin a chapter is given up to “difficulties on theory”: the discussion in the present essay seems slight even when it is remembered how small a space is here available. For Tibia &c. see p. 48.]. It may be objected such perfect organs as eye and ear, could never be formed, in latter less difficulty as gradations more perfect; at first appears monstrous and to «the» end appears difficulty. But think of gradation, even now manifest, (Tibia and Fibula). Everyone will allow if every fossil preserved, gradation infinitely more perfect; for possibility of selection a perfect «?» gradation is required. Different groups of structure, slight gradation in each group, – every analogy renders it probable that intermediate forms have existed. Be it remembered what strange metamorphoses; part of eye, not directly connected with vision, might come to be [thus used] gradually worked in for this end, – swimming bladder by gradation of structure is admitted to belong to the ear system, – rattlesnake. [Woodpecker best adapted to climb.] In some cases gradation not possible, – as vertebræ, – actually vary in domestic animals, – less difficult if growth followed. Looking to whole animals, a bat formed not for flight[85 - This may be interpreted “The general structure of a bat is the same as that of non-flying mammals.”]. Suppose we had flying fish[86 - That is truly winged fish.] and not one of our now called flying fish preserved, who would have guessed intermediate habits. Woodpeckers and tree-frogs both live in countries where no trees[87 - The terrestrial woodpecker of S. America formed the subject of a paper by Darwin, Proc. Zool. Soc., 1870. See Life and Letters, vol. iii. p. 153.].

The gradations by which each individual organ has arrived at its present state, and each individual animal with its aggregate of organs has arrived, probably never could be known, and all present great difficulties. I merely wish to show that the proposition is not so monstrous as it at first appears, and that if good reason can be advanced for believing the species have descended from common parents, the difficulty of imagining intermediate forms of structure not sufficient to make one at once reject the theory.

§ III. «On Variation in instincts and other mental attributes.»

The mental powers of different animals in wild and tame state [present still greater difficulties] require a separate section. Be it remembered I have nothing to do with origin of memory, attention, and the different faculties of the mind[88 - The same proviso occurs in the Origin, Ed. i. p. 207, vi. p. 319.], but merely with their differences in each of the great divisions of nature. Disposition, courage, pertinacity «?», suspicion, restlessness, ill-temper, sagacity and «the» reverse unquestionably vary in animals and are inherited (Cuba wildness dogs, rabbits, fear against particular object as man Galapagos[89 - The tameness of the birds in the Galapagos is described in the Journal of Researches (1860), p. 398. Dogs and rabbits are probably mentioned as cases in which the hereditary fear of man has been lost. In the 1844 MS. the author states that the Cuban feral dog shows great natural wildness, even when caught quite young.]). Habits purely corporeal, breeding season &c., time of going to rest &c., vary and are hereditary, like the analogous habits of plants which vary and are inherited. Habits of body, as manner of movement do. and do. Habits, as pointing and setting on certain occasions do. Taste for hunting certain objects and manner of doing so, – sheep-dog. These are shown clearly by crossing and their analogy with true instinct thus shown, – retriever. Do not know objects for which they do it. Lord Brougham’s definition[90 - In the Origin, Ed. i. p. 207, vi. p. 319, he refuses to define instinct. For Lord Brougham’s definition see his Dissertations on Subjects of Science etc., 1839, p. 27.]. Origin partly habit, but the amount necessarily unknown, partly selection. Young pointers pointing stones and sheep – tumbling pigeons – sheep[91 - See James Hogg (the Ettrick Shepherd), Works, 1865, Tales and Sketches, p. 403.] going back to place where born. Instinct aided by reason, as in the taylor-bird[92 - This refers to the tailor-bird making use of manufactured thread supplied to it, instead of thread twisted by itself.]. Taught by parents, cows choosing food, birds singing. Instincts vary in wild state (birds get wilder) often lost[93 - Often lost applies to instinct: birds get wilder is printed in a parenthesis because it was apparently added as an after-thought. Nest without roof refers to the water-ousel omitting to vault its nest when building in a protected situation.]; more perfect, – nest without roof. These facts [only clear way] show how incomprehensibly brain has power of transmitting intellectual operations.

Faculties[94 - In the MS. of 1844 is an interesting discussion on faculty as distinct from instinct.] distinct from true instincts, – finding [way]. It must I think be admitted that habits whether congenital or acquired by practice [sometimes] often become inherited[95 - At this date and for long afterwards the inheritance of acquired characters was assumed to occur.]; instincts, influence, equally with structure, the preservation of animals; therefore selection must, with changing conditions tend to modify the inherited habits of animals. If this be admitted it will be found possible that many of the strangest instincts may be thus acquired. I may observe, without attempting definition, that an inherited habit or trick (trick because may be born) fulfils closely what we mean by instinct. A habit is often performed unconsciously, the strangest habits become associated, do. tricks, going in certain spots &c. &c., even against will, is excited by external agencies, and looks not to the end, – a person playing a pianoforte. If such a habit were transmitted it would make a marvellous instinct. Let us consider some of the most difficult cases of instincts, whether they could be possibly acquired. I do not say probably, for that belongs to our 3rd Part[96 - Part II. is here intended: see the Introduction.], I beg this may be remembered, nor do I mean to attempt to show exact method. I want only to show that whole theory ought not at once to be rejected on this score.

Every instinct must, by my theory, have been acquired gradually by slight changes «illegible» of former instinct, each change being useful to its then species. Shamming death struck me at first as remarkable objection. I found none really sham death[97 - The meaning is that the attitude assumed in shamming is not accurately like that of death.], and that there is gradation; now no one doubts that those insects which do it either more or less, do it for some good, if then any species was led to do it more, and then «?» escaped &c. &c.

Take migratory instincts, faculty distinct from instinct, animals have notion of time, – like savages. Ordinary finding way by memory, but how does savage find way across country, – as incomprehensible to us, as animal to them, – geological changes, – fishes in river, – case of sheep in Spain[98 - This refers to the transandantes sheep mentioned in the MS. of 1844, as having acquired a migratory instinct.]. Architectural instincts, – a manufacturer’s employee in making single articles extraordinary skill, – often said seem to make it almost «illegible», child born with such a notion of playing[99 - In the Origin, Ed. i. p. 209, vi. p. 321, Mozart’s pseudo-instinctive skill in piano-playing is mentioned. See Phil. Trans., 1770, p. 54.], – we can fancy tailoring acquired in same perfection, – mixture of reason, – water-ouzel, – taylor-bird, – gradation of simple nest to most complicated.

Bees again, distinction of faculty, – how they make a hexagon, – Waterhouse’s theory[100 - In the discussion on bees’ cells, Origin, Ed. i. p. 225, vi. p. 343, the author acknowledges that his theory originated in Waterhouse’s observations.], – the impulse to use whatever faculty they possess, – the taylor-bird has the faculty of sewing with beak, instinct impels him to do it.

Last case of parent feeding young with different food (take case of Galapagos birds, gradation from Hawfinch to Sylvia) selection and habit might lead old birds to vary taste «?» and form, leaving their instinct of feeding their young with same food[101 - The hawfinch-and Sylvia-types are figured in the Journal of Researches, p. 379. The discussion of change of form in relation to change of instinct is not clear, and I find it impossible to suggest a paraphrase.], – or I see no difficulty in parents being forced or induced to vary the food brought, and selection adapting the young ones to it, and thus by degree any amount of diversity might be arrived at. Although we can never hope to see the course revealed by which different instincts have been acquired, for we have only present animals (not well known) to judge of the course of gradation, yet once grant the principle of habits, whether congenital or acquired by experience, being inherited and I can see no limit to the [amount of variation] extraordinariness «?» of the habits thus acquired.

Summing up this Division. If variation be admitted to occur occasionally in some wild animals, and how can we doubt it, when we see [all] thousands «of» organisms, for whatever use taken by man, do vary. If we admit such variations tend to be hereditary, and how can we doubt it when we «remember» resemblances of features and character, – disease and monstrosities inherited and endless races produced (1200 cabbages). If we admit selection is steadily at work, and who will doubt it, when he considers amount of food on an average fixed and reproductive powers act in geometrical ratio. If we admit that external conditions vary, as all geology proclaims, they have done and are now doing, – then, if no law of nature be opposed, there must occasionally be formed races, [slightly] differing from the parent races. So then any such law[102 - I should interpret this obscure sentence as follows, “No such opposing law is known, but in all works on the subject a law is (in flat contradiction to all known facts) assumed to limit the possible amount of variation.” In the Origin, the author never limits the power of variation, as far as I know.], none is known, but in all works it is assumed, in «?» flat contradiction to all known facts, that the amount of possible variation is soon acquired. Are not all the most varied species, the oldest domesticated: who «would» think that horses or corn could be produced? Take dahlia and potato, who will pretend in 5000 years[103 - In Var. under Dom. Ed. 2, ii. p. 263, the Dahlia is described as showing sensitiveness to conditions in 1841. All the varieties of the Dahlia are said to have arisen since 1804 (ibid. i. p. 393).] «that great changes might not be effected»: perfectly adapted to conditions and then again brought into varying conditions. Think what has been done in few last years, look at pigeons, and cattle. With the amount of food man can produce he may have arrived at limit of fatness or size, or thickness of wool «?», but these are the most trivial points, but even in these I conclude it is impossible to say we know the limit of variation. And therefore with the [adapting] selecting power of nature, infinitely wise compared to those of man, «I conclude» that it is impossible to say we know the limit of races, which would be true «to their» kind; if of different constitutions would probably be infertile one with another, and which might be adapted in the most singular and admirable manner, according to their wants, to external nature and to other surrounding organisms, – such races would be species. But is there any evidence «that» species «have» been thus produced, this is a question wholly independent of all previous points, and which on examination of the kingdom of nature «we» ought to answer one way or another.

PART II[104 - In the original MS. the heading is: Part III.; but Part II. is clearly intended; for details see the Introduction. I have not been able to discover where § IV. ends and § V. begins.]

§§ IV. & V. «On the evidence from Geology.»