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- More Letters of Charles Darwin Volume I - 60/99 -

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 constitutional vigour of the hybrids exactly counterbalances their imperfect adaptations to conditions, there will be in the second generation, besides these three classes, 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 succumb, 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 accumulated 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).)

P.S.--In answer to the objection as to the unequal sterility of reciprocal crosses ("Variation, etc." Volume II., page 186) I reply that, as far as it went, the sterility of one cross would be advantageous even if the other cross was fertile: and just as characters now co-ordinated may have been separately accumulated by Natural Selection, so the reciprocal crosses may have become sterile one at a time.

LETTER 212. TO A.R. WALLACE. 4, Chester Place, March 17th, 1868.

(212/1. Mr. Darwin had already written a short note to Mr. Wallace expressing a general dissent from his view.)

I do not feel that I shall grapple with the sterility argument till my return home; I have tried once or twice, and it has made my stomach feel as if it had been placed in a vice. Your paper has driven three of my children half mad--one sat up till 12 o'clock over it. My second son, the mathematician, thinks that you have omitted one almost inevitable deduction which apparently would modify the result. He has written out what he thinks, but I have not tried fully to understand him. I suppose that you do not care enough about the subject to like to see what he has written.

LETTER 212A. A.R. WALLACE TO CHARLES DARWIN. Hurstpierpoint, March, 24th [1868].

I return your son's notes with my notes on them. Without going into any details, is not this a strong general argument?

1. A species varies occasionally in two directions, but owing to their free intercrossing the varieties never increase.

2. A change of conditions occurs which threatens the existence of the species; but the two varieties are adapted to the changing conditions, and if accumulated will form two new species adapted to the new conditions.

3. Free crossing, however, renders this impossible, and so the species is in danger of extinction.

4. If sterility would be induced, then the pure races would increase more rapidly, and replace the old species.

5. It is admitted that partial sterility between varieties does occasionally occur. It is admitted [that] the degree of this sterility varies; is it not probable that Natural Selection can accumulate these variations, and thus save the species? If Natural Selection can NOT do this, how do species ever arise, except when a variety is isolated?

Closely allied species in distinct countries being sterile is no difficulty; for either they diverged from a common ancestor in contact, and Natural Selection increased the sterility, or they were isolated, and have varied since: in which case they have been for ages influenced by distinct conditions which may well produce sterility.

If the difficulty of grafting was as great as the difficulty of crossing, and as regular, I admit it would be a most serious objection. But it is not. I believe many distinct species can be grafted, while others less distinct cannot. The regularity with which natural species are sterile together, even when very much alike, I think is an argument in favour of the sterility having been generally produced by Natural Selection for the good of the species.

The other difficulty, of unequal sterility of reciprocal crosses, seems none to me; for it is a step to more complete sterility, and as such would be increased by selection.

LETTER 213. TO A.R. WALLACE. Down, April 6th [1868].

I have been considering the terrible problem. Let me first say that no man could have more earnestly wished for the success of Natural Selection in regard to sterility than I did; and when I considered a general statement (as in your last note) I always felt sure it could be worked out, but always failed in detail. The cause being, as I believe, that Natural Selection cannot effect what is not good for the individual, including in this term a social community. It would take a volume to discuss all the points, and nothing is so humiliating to me as to agree with a man like you (or Hooker) on the premises and disagree about the result.

I agree with my son's argument and not with the rejoinder. The cause of our difference, I think, is that I look at the number of offspring as an important element (all circumstances remaining the same) in keeping up the average number of individuals within any area. I do not believe that the amount of food by any means is the sole determining cause of number. Lessened fertility is equivalent to a new source of destruction. I believe if in one district a species produced from any cause fewer young, the deficiency would be supplied from surrounding districts. This applies to your Paragraph 5. (213/1. See Letter 211.) If the species produced fewer young from any cause in every district, it would become extinct unless its fertility were augmented through Natural Selection (see H. Spencer).

I demur to probability and almost to possibility of Paragraph 1., as you start with two forms within the same area, which are not mutually sterile, and which yet have supplanted the parent-form.

(Paragraph 6.) I know of no ghost of a fact supporting belief that disinclination to cross accompanies sterility. It cannot hold with plants, or the lower fixed aquatic animals. I saw clearly what an immense aid this would be, but gave it up. Disinclination to cross seems to have been independently acquired, probably by Natural Selection; and I do not see why it would not have sufficed to have prevented incipient species from blending to have simply increased sexual disinclination to cross.

(Paragraph 11.) I demur to a certain extent to amount of sterility and structural dissimilarity necessarily going together, except indirectly and by no means strictly. Look at vars. of pigeons, fowls, and cabbages.

I overlooked the advantage of the half-sterility of reciprocal crosses; yet, perhaps from novelty, I do not feel inclined to admit probability of Natural Selection having done its work so queerly.

I will not discuss the second case of utter sterility, but your assumptions in Paragraph 13 seem to me much too complicated. I cannot believe so universal an attribute as utter sterility between remote species was acquired in so complex a manner. I do not agree with your rejoinder on grafting: I fully admit that it is not so closely restricted as crossing, but this does not seem to me to weaken the case as one of analogy. The incapacity of grafting is likewise an invariable attribute of plants sufficiently remote from each other, and sometimes of plants pretty closely allied.

The difficulty of increasing the sterility through Natural Selection of two already sterile species seems to me best brought home by considering an actual case. The cowslip and primrose are moderately sterile, yet occasionally produce hybrids. Now these hybrids, two or three or a dozen in a whole parish, occupy ground which might have been occupied by either pure species, and no doubt the latter suffer to this small extent. But can you conceive that any individual plants of the primrose and cowslip which happened to be mutually rather more sterile (i.e. which, when crossed, yielded a few less seed) than usual, would profit to such a degree as to increase in number to the ultimate exclusion of the present primrose and cowslip? I cannot.

My son, I am sorry to say, cannot see the full force of your rejoinder in regard to second head of continually augmented sterility. You speak in this rejoinder, and in Paragraph 5, of all the individuals becoming in some slight degree sterile in certain districts: if you were to admit that by continued exposure to these same conditions the sterility would inevitably increase, there would be no need of Natural Selection. But I suspect that the sterility is not caused so much by any particular conditions as by long habituation to conditions of any kind. To speak according to pangenesis, the gemmules of hybrids are not injured, for hybrids propagate freely by buds; but their reproductive organs are somehow affected, so that they cannot accumulate the proper gemmules, in nearly the same manner as the

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