Hildebrand"s paper is in the "Bot. Zeitung," 1868: the substance is given in "Variation of Animals and Plants," Edition II., Volume I., page 420.), for I believe it to be a most important observation. I trust that you will publish full details on this subject and on the direct action of pollen (206/2. See Prof. Hildebrand, "Bot. Zeitung," 1868, and "Variation of Animals and Plants," Edition II., Volume I., page 430.
A yellow-grained maize was fertilised with pollen from a brown-grained one; the result was that ears were produced bearing both yellow and dark-coloured grains.): I hope that you will be so kind as to send me a copy of your paper. If I had succeeded in making a graft-hybrid of the potato, I had intended to raise seedlings from the graft-hybrid and from the two parent-forms (excluding insects) and carefully compare the offspring. This, however, would be difficult on account of the sterility and variability of the potato. When in the course of a few months you receive my second volume (206/3. This sentence may be paraphrased--"When you receive my book and read the second volume."), you will see why I think these two subjects so important. They have led me to form a hypothesis on the various forms of reproduction, development, inheritance, etc., which hypothesis, I believe, will ultimately be accepted, though how it will be now received I am very doubtful.
Once again I congratulate you on your success.
LETTER 207. TO J.D. HOOKER. Down, January 6th [1868].
Many thanks about names of plants, synonyms, and male flowers--all that I wanted.
I have been glad to see Watson"s letter, and am sorry he is a renegade about Natural Selection. It is, as you say, characteristic, with the final fling at you.
His difficulty about the difference between the two genera of St. Helena Umbellifers is exactly the same as what Nageli has urged in an able pamphlet (207/1. "Ueber Entstehung und Begriff der naturhist. Art."
"Sitz. der K. Bayer. Akad. Der Wiss. zu Munchen," 1865. Some of Nageli"s points are discussed in the "Origin," Edition V., page 151.), and who in consequence maintains that there is some unknown innate tendency to progression in all organisms. I said in a letter to him that of course I could not in the least explain such cases; but that they did not seem to me of overwhelming force, as long as we are quite ignorant of the meaning of such structures, whether they are of any service to the plants, or inevitable consequences of modifications in other parts.
I cannot understand what Watson means by the "counter-balance in nature"
to divergent variation. There is the counterbalance of crossing, of which my present work daily leads me to see more and more the efficiency; but I suppose he means something very different. Further, I believe variation to be divergent solely because diversified forms can best subsist. But you will think me a bore.
I enclose half a letter from F. Muller (which please return) for the chance of your liking to see it; though I have doubted much about sending it, as you are so overworked. I imagine the Solanum-like flower is curious.
I heard yesterday to my joy that Dr. Hildebrand has been experimenting on the direct action of pollen on the mother-plant with success. He has also succeeded in making a true graft-hybrid between two varieties of potatoes, in which I failed. I look at this as splendid for pangenesis, as being strong evidence that bud-reproduction and seminal reproduction do not essentially differ.
My book is horribly delayed, owing to the accursed index-maker. (207/2.
Darwin thoroughly appreciated the good work put into the index of "The Variation of Animals and Plants.") I have almost forgotten it!
LETTER 208. TO T.H. HUXLEY. Down, January 30th [1868].
Most sincere thanks for your kind congratulations. I never received a note from you in my life without pleasure; but whether this will be so after you have read pangenesis (208/1. In Volume II. of "Animals and Plants, 1868.), I am very doubtful. Oh Lord, what a blowing up I may receive! I write now partly to say that you must not think of looking at my book till the summer, when I hope you will read pangenesis, for I care for your opinion on such a subject more than for that of any other man in Europe. You are so terribly sharp-sighted and so confoundedly honest! But to the day of my death I will always maintain that you have been too sharp-sighted on hybridism; and the chapter on the subject in my book I should like you to read: not that, as I fear, it will produce any good effect, and be hanged to you.
I rejoice that your children are all pretty well. Give Mrs. Huxley the enclosed (208/2. Queries on Expression.), and ask her to look out when one of her children is struggling and just going to burst out crying. A dear young lady near here plagued a very young child for my sake, till it cried, and saw the eyebrows for a second or two beautifully oblique, just before the torrent of tears began.
The sympathy of all our friends about George"s success (it is the young Herald) (208/3. His son George was Second Wrangler in 1868; as a boy he was an enthusiast in heraldry.) has been a wonderful pleasure to us. George has not slaved himself, which makes his success the more satisfactory. Farewell, my dear Huxley, and do not kill yourself with work.
(209/1. The following group of letters deals with the problem of the causes of the sterility of hybrids. Mr. Darwin"s final view is given in the "Origin," sixth edition (page 384, edition 1900). He acknowledges that it would be advantageous to two incipient species, if by physiological isolation due to mutual sterility, they could be kept from blending: but he continues, "After mature reflection it seems to me that this could not have been effected through Natural Selection." And finally he concludes (page 386):--
"But it would be superfluous to discuss this question in detail; for with plants we have conclusive evidence that the sterility of crossed species must be due to some principle quite independent of Natural Selection. Both Gartner and Kolreuter have proved that in genera including numerous species, a series can be formed from species which when crossed yield fewer and fewer seeds, to species which never produce a single seed, but yet are affected by the pollen of certain other species, for the germen swells. It is here manifestly impossible to select the more sterile individuals, which have already ceased to yield seeds; so that this acme of sterility, when the germen alone is affected, cannot have been gained through selection; and from the laws governing the various grades of sterility being so uniform throughout the animal and vegetable kingdoms, we may infer that the cause, whatever it may be, is the same or nearly the same in all cases."
Mr. Wallace, on the other hand, still adheres to his view: see his "Darwinism," 1889, page 174, and for a more recent statement see page 292, note 1, Letter 211, and page 299.
The discussion of 1868 began with a letter from Mr. Wallace, written towards the end of February, giving his opinion on the "Variation of Animals and Plants;" the discussion on the sterility of hybrids is at page 185, Volume II., of the first edition.)
LETTER 209. A.R. WALLACE TO CHARLES DARWIN. February 1868.
The only parts I have yet met with where I somewhat differ from your views, are in the chapter on the causes of variability, in which I think several of your arguments are unsound: but this is too long a subject to go into now. Also, I do not see your objection to sterility between allied species having been aided by Natural Selection. It appears to me that, given a differentiation of a species into two forms, each of which was adapted to a special sphere of existence, every slight degree of sterility would be a positive advantage, not to the individuals who were sterile, but to each form. If you work it out, and suppose the two incipient species a...b to be divided into two groups, one of which contains those which are fertile when the two are crossed, the other being slightly sterile, you will find that the latter will certainly supplant the former in the struggle for existence; remembering that you have shown that in such a cross the offspring would be more vigorous than the pure breed, and therefore would certainly soon supplant them, and as these would not be so well adapted to any special sphere of existence as the pure species a and b, they would certainly in their turn give way to a and b.
LETTER 210. TO A.R. WALLACE. February 27th [1868].
I shall be very glad to hear, at some future day, your criticisms on the "causes of variability." Indeed, I feel sure that I am right about sterility and Natural Selection. Two of my grown-up children who are acute reasoners have two or three times at intervals tried to prove me wrong; and when your letter came they had another try, but ended by coming back to my side. I do not quite understand your case, and we think that a word or two is misplaced. I wish some time you would consider the case under the following point of view. If sterility is caused or acc.u.mulated through Natural Selection, then, as every degree exists up to absolute barrenness, Natural Selection must have the power of increasing it. Now take two species A and B, and a.s.sume that they are (by any means) half-sterile, i.e., produce half the full number of offspring. Now try and make (by Natural Selection) A and B absolutely sterile when crossed, and you will find how difficult it is. I grant, indeed it is certain, that the degree of the sterility of the individuals of A and B will vary; but any such extra-sterile individuals of, we will say A, if they should hereafter breed with other individuals of A, will bequeath no advantage to their progeny, by which these families will tend to increase in number over other families of A, which are not more sterile when crossed with B. But I do not know that I have made this any clearer than in the chapter in my book. It is a most difficult bit of reasoning, which I have gone over and over again on paper with diagrams. (210/1. This letter appeared in "Life and Letters,"
III., page 80.)
LETTER 211. A.R. WALLACE TO CHARLES DARWIN. March 1st, 1868.
I beg to enclose what appears to me a demonstration on your own principles, that Natural Selection could produce sterility of hybrids.
If it does not convince you, I shall be glad if you will point out where the fallacy lies. I have taken the two cases of a slight sterility overcoming perfect fertility, and of a perfect sterility overcoming a partial fertility,--the beginning and end of the process. You admit that variations in fertility and sterility occur, and I think you will also admit that if I demonstrate that a considerable amount of sterility would be advantageous to a variety, that is sufficient proof that the slightest variation in that direction would be useful also, and would go on acc.u.mulating.
1. Let there be a species which has varied into two forms, each adapted to existing conditions (211/1. "Existing conditions," means of course new conditions which have now come into existence. And the "two" being both better adapted than the parent form, means that they are better adapted each to a special environment in the same area--as one to damp, another to dry places; one to woods, another to open grounds, etc., etc., as Darwin had already explained. A.R.W. (1899).) better than the parent form, which they supplant.
2. If these two forms, which are supposed to co-exist in the same district, do not intercross, Natural Selection will acc.u.mulate favourable variations, till they become sufficiently well adapted to their conditions of life and form two allied species.
3. But if these two forms freely intercross with each other and produce hybrids which are also quite fertile inter se, then the formation of the two distinct races or species will be r.e.t.a.r.ded or perhaps entirely prevented; for the offspring of the crossed unions will be more vigorous owing to the cross, although less adapted to their conditions of life than either of the pure breeds. (211/2. After "pure breeds," add "because less specialised." A.R.W. (1899).)
4. Now let a partial sterility of some individuals of these two forms arise when they intercross; and as this would probably be due to some special conditions of life, we may fairly suppose it to arise in some definite portion of the area occupied by the two forms.
5. The result is that in this area hybrids will not increase so rapidly as before; and as by the terms of the problem the two pure forms are better suited to the conditions of life than the hybrids, they will tend to supplant the latter altogether whenever the struggle for existence becomes severe.
6. We may fairly suppose, also, that as soon as any sterility appears under natural conditions, it will be accompanied by some disinclination to cross-unions; and this will further diminish the production of hybrids.
7. In the other part of the area, however, where hybridism occurs unchecked, hybrids of various degrees will soon far outnumber the parent or pure form.
8. The first result, then, of a partial sterility of crosses appearing in one part of the area occupied by the two forms, will be, that the GREAT MAJORITY of the individuals will there consist of the pure forms only, while in the rest of the area these will be in a minority,--which is the same as saying, that the new sterile or physiological variety of the two forms will be better suited to the conditions of existence than the remaining portion which has not varied physiologically.
9. But when the struggle for existence becomes severe, that variety which is best adapted to the conditions of existence always supplants that which is imperfectly adapted; therefore by Natural Selection the sterile varieties of the two forms will become established as the only ones.
10. Now let a fresh series of variations in the amount of sterility and in the disinclination to crossed unions occur,--also in certain parts of the area: exactly the same result must recur, and the progeny of this new physiological variety again in time occupy the whole area.
11. There is yet another consideration that supports this view. It seems probable that the variations in amount of sterility would to some extent concur with and perhaps depend upon the structural variations; so that just in proportion as the two forms diverged and became better adapted to the conditions of existence, their sterility would increase. If this were the case, then Natural Selection would act with double strength, and those varieties which were better adapted to survive both structurally and physiologically, would certainly do so. (211/3. The preceding eleven paragraphs are substantially but not verbally identical with the statement of the argument in Mr. Wallace"s "Darwinism," 1889.
Pages 179, 180, note 1.)
12. Let us now consider the more difficult case of two allied species A, B, in the same area, half the individuals of each (As, Bs) being absolutely sterile, the other half (Af, Bf) being partially fertile: will As, Bs ultimately exterminate Af, Bf?
13. To avoid complication, it must be granted, that between As and Bs no cross-unions take place, while between Af and Bf cross-unions are as frequent as direct unions, though much less fertile. We must also leave out of consideration crosses between As and Af, Bs and Bf, with their various approaches to sterility, as I believe they will not affect the final result, although they will greatly complicate the problem.
14. In the first generation there will result: 1st, The pure progeny of As and Bs; 2nd, The pure progeny of Af and of Bf; and 3rd, The hybrid progeny of Af, Bf.
15. Supposing that, in ordinary years, the increased const.i.tutional vigour of the hybrids exactly counterbalances their imperfect adaptations to conditions, there will be in the second generation, besides these three cla.s.ses, hybrids of the second degree between the first hybrids and Af and Bf respectively. In succeeding generations there will be hybrids of all degrees, varying between the first hybrids and the almost pure types of Af and Bf.
16. Now, if at first the number of individuals of As, Bs, Af and Bf were equal, and year after year the total number continues stationary, I think it can be proved that, while half will be the pure progeny of As and Bs, the other half will become more and more hybridised, until the whole will be hybrids of various degrees.
17. Now, this hybrid and somewhat intermediate race cannot be so well adapted to the conditions of life as the two pure species, which have been formed by the minute adaptation to conditions through Natural Selection; therefore, in a severe struggle for existence, the hybrids must succ.u.mb, especially as, by hypothesis, their fertility would not be so great as that of the two pure species.
18. If we were to take into consideration the unions of As with Af and Bs with Bf, the results would become very complicated, but it must still lead to there being a number of pure forms entirely derived from As and Bs, and of hybrid forms mainly derived from Af and Bf; and the result of the struggle of these two sets of individuals cannot be doubtful.
19. If these arguments are sound, it follows that sterility may be acc.u.mulated and increased, and finally made complete by Natural Selection, whether the sterile varieties originate together in a definite portion of the area occupied by the two species, or occur scattered over the whole area. (211/4. The first part of this discussion should be considered alone, as it is both more simple and more important. I now believe that the utility, and therefore the cause of sterility between species, is during the process of differentiation.
When species are fully formed, the occasional occurrence of hybrids is of comparatively small importance, and can never be a danger to the existence of the species. A.R.W. (1899).)