
The Foundations of the Origin of Species
I have said that man selects in full-life, so would it be in Nature. In struggle of existence, it matters nothing to a feline animal, whether kitten eminently feline, as long as it sucks. Therefore natural selection would act equally well on character which was fully «developed» only in full age. Selection could tend to alter no character in fœtus, (except relation to mother) it would alter less in young state (putting on one side larva condition) but alter every part in full-grown condition. Look to a fœtus and its parent, and again after ages fœtus and its «i. e. the above mentioned parents» descendant; the parent more variable «?» than fœtus, which explains all.
165
Some of these examples occur in Origin, Ed. i. pp. 450-51, vi. pp. 619-20.
166
The two following sentences are written, one down the margin, the other across the page. “Abortive organs eminently useful in classification. Embryonic state of organs. Rudiments of organs.”
167
I imagine the meaning to be that abortive organs are specific characters in contrast to monstrosities.
168
Minute hanging horns are mentioned in the Origin, Ed. i. p. 454, vi. p. 625, as occurring in hornless breeds of cattle.
169
Linum flavum is dimorphic: thyme gynodiæcious. It is not clear what point is referred to under Geranium pyrenaicum.
170
The author’s work on duck’s wings &c. is in Var. under Dom., Ed. 2, i. p. 299.
171
The words vis medicatrix are inserted after “useless,” apparently as a memorandum.
172
In the male florets of certain Compositæ the style functions merely as a piston for forcing out the pollen.
173
«On the back of the page is the following.» If abortive organs are a trace preserved by hereditary tendency, of organ in ancestor of use, we can at once see why important in natural classification, also why more plain in young animal because, as in last section, the selection has altered the old animal most. I repeat, these wondrous facts, of parts created for no use in past and present time, all can by my theory receive simple explanation; or they receive none and we must be content with some such empty metaphor, as that of De Candolle, who compares creation to a well covered table, and says abortive organs may be compared to the dishes (some should be empty) placed symmetrically!
174
The author doubtless meant that the complex relationships between organisms can be roughly represented by a net in which the knots stand for species.
175
Between the lines occurs: – “one «?» form be lost.”
176
The original sentence is here broken up by the insertion of: – “out of the dust of Java, Sumatra, these «?» allied to past and present age and «illegible», with the stamp of inutility in some of their organs and conversion in others.”
177
Between the lines occur the words: – “Species vary according to same general laws as varieties; they cross according to same laws.”
178
“A cross with a bull-dog has affected for many generations the courage and obstinacy of greyhounds,” Origin, Ed. i. p. 214, vi. p. 327.
179
The simile of the savage and the ship occurs in the Origin, Ed. i. p. 485, vi. p. 665.
180
In the Origin, Ed. i. p. 486, vi. p. 665, the author speaks of the “summing up of many contrivances”: I have therefore introduced the above words which make the passage clearer. In the Origin the comparison is with “a great mechanical invention,” – not with a work of art.
181
See a similar passage in the Origin, Ed. i. p. 487, vi. p. 667.
182
See the Origin, Ed. i. p. 488, vi. p. 668.
183
The following discussion, together with some memoranda are on the last page of the MS. “The supposed creative spirit does not create either number or kind which «are» from analogy adapted to site (viz. New Zealand): it does not keep them all permanently adapted to any country, – it works on spots or areas of creation, – it is not persistent for great periods, – it creates forms of same groups in same regions, with no physical similarity, – it creates, on islands or mountain summits, species allied to the neighbouring ones, and not allied to alpine nature as shown in other mountain summits – even different on different island of similarly constituted archipelago, not created on two points: never mammifers created on small isolated island; nor number of organisms adapted to locality: its power seems influenced or related to the range of other species wholly distinct of the same genus, – it does not equally effect, in amount of difference, all the groups of the same class.”
184
This passage is the ancestor of the concluding words in the first edition of the Origin of Species which have remained substantially unchanged throughout subsequent editions, “There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.” In the 2nd edition “by the Creator” is introduced after “originally breathed.”
185
Compare the Origin, Ed. i. p. 481, vi. p. 659, “The difficulty is the same as that felt by so many geologists, when Lyell first insisted that long lines of inland cliffs had been formed, and great valleys excavated, by the slow action of the coast-waves.”
186
The cumulative effect of domestication is insisted on in the Origin, see e. g. Origin, Ed. i. p. 7, vi. p. 8.
187
This type of variation passes into what he describes as the direct effect of conditions. Since they are due to causes acting during the adult life of the organism they might be called individual variations, but he uses this term for congenital variations, e. g. the differences discoverable in plants raised from seeds of the same pod (Origin, Ed. i. p. 45, vi. p. 53).
188
«It is not clear where the following note is meant to come»: Case of Orchis, – most remarkable as not long cultivated by seminal propagation. Case of varieties which soon acquire, like Ægilops and Carrot (and Maize) a certain general character and then go on varying.
189
Here, as in the MS. of 1842, the author is inclined to minimise the variation occurring in nature.
190
This is more strongly stated than in the Origin, Ed. i. p. 30.
191
See Origin, Ed. i. p. 13.
192
Origin, Ed. i. p. 86, vi. p. 105.
193
It is interesting to find that though the author, like his contemporaries, believed in the inheritance of acquired characters, he excluded the case of mutilation.
194
This corresponds to Origin, Ed. i. p. 10, vi. p. 9.
195
Origin, Ed. i. p. 8, vi. p. 10.
196
For plasticity see Origin, Ed. i. pp. 12, 132.
197
Var. under Dom., Ed. ii. I. p. 393.
198
Selection is here used in the sense of isolation, rather than as implying the summation of small differences. Professor Henslow in his Heredity of Acquired Characters in Plants, 1908, p. 2, quotes from Darwin’s Var. under Dom., Ed. i. II. p. 271, a passage in which the author, speaking of the direct action of conditions, says: – “A new sub-variety would thus be produced without the aid of selection.” Darwin certainly did not mean to imply that such varieties are freed from the action of natural selection, but merely that a new form may appear without summation of new characters. Professor Henslow is apparently unaware that the above passage is omitted in the second edition of Var. under Dom., II. p. 260.
199
See the Essay of 1842, p. 3.
200
See Origin, Ed. i. p. 33, vi. p. 38. The evidence is given in the present Essay rather more fully than in the Origin.
201
Journal of Researches, Ed. 1860, p. 214. “Doggies catch otters, old women no.”
202
The effects of crossing is much more strongly stated here than in the Origin. See Ed. i. p. 20, vi. p. 23, where indeed the opposite point of view is given. His change of opinion may be due to his work on pigeons. The whole of the discussion on crossing corresponds to Chapter VIII of the Origin, Ed. i. rather than to anything in the earlier part of the book.
203
The parallelism between the effects of a cross and the effects of conditions is given from a different point of view in the Origin, Ed. i. p. 266, vi. p. 391. See the experimental evidence for this important principle in the author’s work on Cross and Self-Fertilisation. Professor Bateson has suggested that the experiments should be repeated with gametically pure plants.
204
The so-called Knight-Darwin Law is often misunderstood. See Goebel in Darwin and Modern Science, 1909, p. 419; also F. Darwin, Nature, Oct. 27, 1898.
205
Pallas’ theory is discussed in the Origin, Ed. i. pp. 253, 254, vi. p. 374.
206
See Darwin’s paper on the fertility of hybrids from the common and Chinese goose in Nature, Jan. 1, 1880.
207
Origin, Ed. i. p. 19, vi. p. 22.
208
Var. under Dom., Ed. ii. vol. II. p. 211.
209
This discussion corresponds to the Origin, Ed. i. pp. 11 and 143, vi. pp. 13 and 177.
210
See Origin, Ed. i. p. 7, vi. p. 7.
211
«Note in the original.» “Isidore G. St Hilaire insists that breeding in captivity essential element. Schleiden on alkalies. «See Var. under Dom., Ed. ii. vol. II. p. 244, note 10.» What is it in domestication which causes variation?”
212
«Note in the original.» “It appears that slight changes of condition «are» good for health; that more change affects the generative system, so that variation results in the offspring; that still more change checks or destroys fertility not of the offspring.” Compare the Origin, Ed. i. p. 9, vi. p. 11. What the meaning of “not of the offspring” may be is not clear.
213
In the Origin, Ed. i. p. 41, vi. p. 46 the question is differently treated; it is pointed out that a large stock of individuals gives a better chance of available variations occurring. Darwin quotes from Marshall that sheep in small lots can never be improved. This comes from Marshall’s Review of the Reports to the Board of Agriculture, 1808, p. 406. In this Essay the name Marshall occurs in the margin. Probably this refers to loc. cit. p. 200, where unshepherded sheep in many parts of England are said to be similar owing to mixed breeding not being avoided.
214
See Origin, Ed. i. p. 8, vi. p. 8.
215
See Origin, Ed. i. p. 42, vi. p. 48.
216
«Note in the original.» There are white peacocks.
217
«Note in the original.» There are varieties of asparagus.
218
In Chapter II of the first edition of the Origin Darwin insists rather on the presence of variability in a state of nature; see, for instance, p. 45, Ed. vi. p. 53, “I am convinced that the most experienced naturalist would be surprised at the number of the cases of variability … which he could collect on good authority, as I have collected, during a course of years.”
219
See Origin, Ed. i. p. 44, vi. p. 52.
220
«Note in the original.» Here discuss what is a species, sterility can most rarely be told when crossed. – Descent from common stock.
221
«Note in the original.» Give only rule: chain of intermediate forms, and analogy; this important. Every Naturalist at first when he gets hold of new variable type is quite puzzled to know what to think species and what variations.
222
The author had not at this time the knowledge of the meaning of dimorphism.
223
«Note in original.» Compare feathered heads in very different birds with spines in Echidna and Hedgehog. «In Variation under Domestication, Ed. ii. vol. II. p. 317, Darwin calls attention to laced and frizzled breeds occurring in both fowls and pigeons. In the same way a peculiar form of covering occurs in Echidna and the hedgehog.»
Plants under very different climate not varying. Digitalis shows jumps «?» in variation, like Laburnum and Orchis case – in fact hostile cases. Variability of sexual characters alike in domestic and wild.
224
A corresponding passage occurs in Origin, Ed. i. p. 83, vi. p. 101, where however Nature takes the place of the selecting Being.
225
The mistletoe is used as an illustration in Origin, Ed. i. p. 3, vi. p. 3, but with less detail.
226
«Note in original.» The selection, in cases where adult lives only few hours as Ephemera, must fall on larva – curious speculation of the effect «which» changes in it would bring in parent.
227
This section forms part of the joint paper by Darwin and Wallace read before the Linnean Society on July 1, 1858.
228
Occurs in Origin, Ed. i. p. 64, vi. p. 79.
229
Corresponds approximately with Origin, Ed. i. pp. 64-65, vi. p. 80.
230
This simile occurs in Origin, Ed. i. p. 67, not in the later editions.
231
«Note in the original.» In case like mistletoe, it may be asked why not more species, no other species interferes; answer almost sufficient, same causes which check the multiplication of individuals.
232
See Origin, Ed. i. pp. 104, 292, vi. pp. 127, 429.
233
Recognition of the importance of minute differences in the struggle occurs in the Essay of 1842, p. 8 note 3.
234
See Origin, Ed. i. p. 90, vi. p. 110.
235
These two forms of sexual selection are given in Origin, Ed. i. p. 87, vi. p. 107. The Guiana rock-thrush is given as an example of bloodless competition.
236
«Note in original.» Seals? Pennant about battles of seals.
237
In the Linnean paper of July 1, 1858 the final word is mate: but the context shows that it should be male; it is moreover clearly so written in the MS.
238
In the Origin the author would here have used the word variety.
239
The whole of p. 94 and 15 lines of p. 95 are, in the MS., marked through in pencil with vertical lines, beginning at “Races produced, &c.” and ending with “to these conditions.”
240
See Origin, Ed. i. p. 83, vi. p. 102.
241
In the present Essay there is some evidence that the author attributed more to sports than was afterwards the case: but the above passage points the other way. It must always be remembered that many of the minute differences, now considered small mutations, are the small variations on which Darwin conceived selection to act.
242
See Var. under Dom., Ed. ii. vol. II. p. 230.
243
«Note in the original.» If domestic animals are descended from several species and become fertile inter se, then one can see they gain fertility by becoming adapted to new conditions and certainly domestic animals can withstand changes of climate without loss of fertility in an astonishing manner.
244
See Suchetet, L’Hybridité dans la Nature, Bruxelles, 1888, p. 67. In Var. under Dom., Ed. ii. vol. II. hybrids between the fowl and the pheasant are mentioned. I can give no information on the other cases.
245
Origin, Ed. i. p. 250, vi. p. 370.
246
This was the position of Gärtner and of Kölreuter: see Origin, Ed. i. pp. 246-7, vi. pp. 367-8.
247
«Note in the original.» Yet this seems introductory to the case of the heaths and crocuses above mentioned. «Herbert observed that crocus does not set seed if transplanted before pollination, but that such treatment after pollination has no sterilising effect. (Var. under Dom., Ed. ii. vol. II. p. 148.) On the same page is a mention of the Ericaceæ being subject to contabescence of the anthers. For Crinum see Origin, Ed. i. p. 250: for Rhododenron and Calceolaria see p. 251.»
248
«Note in original.» Animals seem more often made sterile by being taken out of their native condition than plants, and so are more sterile when crossed.
We have one broad fact that sterility in hybrids is not closely related to external difference, and these are what man alone gets by selection.
249
See Var. under Dom., Ed. ii. vol. II. p. 132; for the case of the cheetah see loc cit. p. 133.
250
Var. under Dom., Ed. ii. vol. II. p. 148.
251
Quoted in the Origin, Ed. i. p. 9.
252
See Var. under Dom., Ed. ii. vol. II. p. 147.
253
Var. under Dom., Ed. ii. vol. II. p. 89.
254
See Var. under Dom., Ed. ii. vol. II. p. 147.
255
Origin, Ed. i. p. 267, vi. p. 392. This is the principle experimentally investigated in the author’s Cross-and Self-Fertilisation.
256
Origin, Ed. i. p. 268, vi. p. 398.
257
«Notes in original.» Mere difference of structure no guide to what will or will not cross. First step gained by races keeping apart. «It is not clear where these notes were meant to go.»
258
Origin, Ed. i. p. 272, vi. p. 404.
259
This section seems not to correspond closely with any in the Origin, Ed. i.; in some points it resembles pp. 15, 16, also the section on analogous variation in distinct species, Origin, Ed. i. p. 159, vi. p. 194.
260
The law of compensation is discussed in the Origin, Ed. i. p. 147, vi. p. 182.
261
«Note in original.» Boitard and Corbié on outer edging red in tail of bird, – so bars on wing, white or black or brown, or white edged with black or «illegible»: analogous to marks running through genera but with different colours. Tail coloured in pigeons.
262
«Note in original.» Oxalis and Gentian. «In Gentians blue, yellow and reddish colours occur. In Oxalis yellow, purple, violet and pink.»
263
This section corresponds roughly to that on Hybrids and Mongrels compared independently of their fertility, Origin, Ed. i. p. 272, vi. p. 403. The discussion on Gärtner’s views, given in the Origin, is here wanting. The brief mention of prepotency is common to them both.
264
See Animals and Plants, Ed. ii. vol. I. p. 435. The phenomenon of Telegony, supposed to be established by this and similar cases, is now generally discredited in consequence of Ewart’s experiments.
265
The section on p. 109 is an appendix to the summary.
266
I do not know the authority for this statement.
267
In the Origin no limit is placed to variation as far as I know.
268
«Note in original.» History of pigeons shows increase of peculiarities during last years.
269
Compare an obscure passage in the Essay of 1842, p. 14.
270
«Note in original.» Certainly «two pages in the MS.» ought to be here introduced, viz., difficulty in forming such organ, as eye, by selection. «In the Origin, Ed. i., a chapter on Difficulties on Theory follows that on Laws of Variation, and precedes that on Instinct: this was also the arrangement in the Essay of 1842; whereas in the present Essay Instinct follows Variation and precedes Difficulties.»
271
A similar proviso occurs in the chapter on instinct in Origin, Ed. i. p. 207, vi. p. 319.
272
The discussion occurs later in Chapter VII of the Origin, Ed. i. than in the present Essay, where moreover it is fuller in some respects.
273
In the margin occurs the name of Poeppig. In Var. under Dom., Ed. ii. vol. I. p. 28, the reference to Poeppig on the Cuban dogs contains no mention of the wildness of their offspring.
274
«Note in original.» Several authors.
275
In the margin “Hogg” occurs as authority for this fact. For the reference, see p. 17, note 4.
276
In the Origin, Ed. i., he speaks more decidedly against the belief that instincts are hereditary habits, see for instance pp. 209, 214, Ed. vi. pp. 321, 327. He allows, however, something to habit (p. 216).
277
A suggestion of Hering’s and S. Butler’s views on memory and inheritance. It is not, however, implied that Darwin was inclined to accept these opinions.
278
Lord Brougham’s Dissertations on Subjects of Science, etc., 1839, p. 27.
279
This case is more briefly given in the Origin, Ed. i. p. 213, vi. p. 326. The simile of the butterfly occurs there also.
280
“A little dose, as Pierre Huber expresses it, of judgment or reason, often comes into play.” Origin, Ed. i. p. 208, vi. p. 320.
281
In the margin is written “Retriever killing one bird.” This refers to the cases given in the Descent of Man, 2nd Ed. (in 1 vol.) p. 78, of a retriever being puzzled how to deal with a wounded and a dead bird, killed the former and carried both at once. This was the only known instance of her wilfully injuring game.
282
See Origin, Ed. i. p. 214, vi. p. 327.
283
«Note in original.» Give some definition of instinct, or at least give chief attributes. «In Origin, Ed. i. p. 207, vi. p. 319, Darwin refuses to define instinct.» The term instinct is often used in «a» sense which implies no more than that the animal does the action in question. Faculties and instincts may I think be imperfectly separated. The mole has the faculty of scratching burrows, and the instinct to apply it. The bird of passage has the faculty of finding its way and the instinct to put it in action at certain periods. It can hardly be said to have the faculty of knowing the time, for it can possess no means, without indeed it be some consciousness of passing sensations. Think over all habitual actions and see whether faculties and instincts can be separated. We have faculty of waking in the night, if an instinct prompted us to do something at certain hour of night or day. Savages finding their way. Wrangel’s account – probably a faculty inexplicable by the possessor. There are besides faculties “means,” as conversion of larvæ into neuters and queens. I think all this generally implied, anyhow useful. «This discussion, which does not occur in the Origin, is a first draft of that which follows in the text, p. 123.»