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natural selection。 But I am bound to confess; that; with all my faith in this principle; I should never have anticipated that natural selection could have been efficient in so high a degree; had not the case of these neuter insects convinced me of the fact。 I have; therefore; discussed this case; at some little but wholly insufficient length; in order to show the power of natural selection; and likewise because this is by far the most serious special difficulty; which my theory has encountered。 The case; also; is very interesting; as it proves that with animals; as with plants; any amount of modification in structure can be effected by the accumulation of numerous; slight; and as we must call them accidental; variations; which are in any manner profitable; without exercise or habit having come into play。 For no amount of exercise; or habit; or volition; in the utterly sterile members of a community could possibly have affected the structure or instincts of the fertile members; which alone leave descendants。 I am surprised that no one has advanced this demonstrative case of neuter insects; against the well…known doctrine of Lamarck。 
Summary。 I have endeavoured briefly in this chapter to show that the mental qualities of our domestic animals vary; and that the variations are inherited。 Still more briefly I have attempted to show that instincts vary slightly in a state of nature。 No one will dispute that instincts are of the highest importance to each animal。 Therefore I can see no difficulty; under changing conditions of life; in natural selection accumulating slight modifications of instinct to any extent; in any useful direction。 In some cases habit or use and disuse have probably come into play。 I do not pretend that the facts given in this chapter strengthen in any great degree my theory; but none of the cases of difficulty; to the best of my judgment; annihilate it。 On the other hand; the fact that instincts are not always absolutely perfect and are liable to mistakes; that no instinct has been produced for the exclusive good of other animals; but that each animal takes advantage of the instincts of others; that the canon in natural history; of 'natura non facit saltum' is applicable to instincts as well as to corporeal structure; and is plainly explicable on the foregoing views; but is otherwise inexplicable; all tend to corroborate the theory of natural selection。 
This theory is; also; strengthened by some few other facts in regard to instincts; as by that common case of closely allied; but certainly distinct; species; when inhabiting distant parts of the world and living under considerably different conditions of life; yet often retaining nearly the same instincts。 For instance; we can understand on the principle of inheritance; how it is that the thrush of South America lines its nest with mud; in the same peculiar manner as does our British thrush: how it is that the male wrens (Troglodytes) of North America; build 'cock…nests;' to roost in; like the males of our distinct Kitty…wrens; a habit wholly unlike that of any other known bird。 Finally; it may not be a logical deduction; but to my imagination it is far more satisfactory to look at such instincts as the young cuckoo ejecting its foster…brothers; ants making slaves;  the larvae of ichneumonidae feeding within the live bodies of caterpillars; not as specially endowed or created instincts; but as small consequences of one general law; leading to the advancement of all organic beings; namely; multiply; vary; let the strongest live and the weakest die。  

Chapter 8 … Hybridism

THE view generally entertained by naturalists is that species; when intercrossed; have been specially endowed with the quality of sterility; in order to prevent the confusion of all organic forms。 This view certainly seems at first probable; for species within the same country could hardly have kept distinct had they been capable of crossing freely。 The importance of the fact that hybrids are very generally sterile; has; I think; been much underrated by some late writers。 On the theory of natural selection the case is especially important; inasmuch as the sterility of hybrids could not possibly be of any advantage to them; and therefore could not have been acquired by the continued preservation of successive profitable degrees of sterility。 I hope; however; to be able to show that sterility is not a specially acquired or endowed quality; but is incidental on other acquired differences。 
In treating this subject; two classes of facts; to a large extent fundamentally different; have generally been confounded together; namely; the sterility of two species when first crossed; and the sterility of the hybrids produced from them。 
Pure species have of course their organs of reproduction in a perfect condition; yet when intercrossed they produce either few or no offspring。 Hybrids; on the other hand; have their reproductive organs functionally impotent; as may be clearly seen in the state of the male element in both plants and animals; though the organs themselves are perfect in structure; as far as the microscope reveals。 In the first case the two sexual elements which go to form the embryo are perfect; in the second case they are either not at all developed; or are imperfectly developed。 This distinction is important; when the cause of the sterility; which is common to the two cases; has to be considered。 The distinction has probably been slurred over; owing to the sterility in both cases being looked on as a special endowment; beyond the province of our reasoning powers。 
The fertility of varieties; that is of the forms known or believed to have descended from common parents; when intercrossed; and likewise the fertility of their mongrel offspring; is; on my theory; of equal importance with the sterility of species; for it seems to make a broad and clear distinction between varieties and species。 
First; for the sterility of species when crossed and of their hybrid offspring。 It is impossible to study the several memoirs and works of those two conscientious and admirable observers; K?lreuter and G?rtner; who almost devoted their lives to this subject; without being deeply impressed with the high generality of some degree of sterility。 K?lreuter makes the rule universal; but then he cuts the knot; for in ten cases in which he found two forms; considered by most authors as distinct species; quite fertile together; he unhesitatingly ranks them as varieties。 G?rtner; also; makes the rule equally universal; and he disputes the entire fertility of K?lreuter's ten cases。 But in these and in many other cases; G?rtner is obliged carefully to count the seeds; in order to show that there is any degree of sterility。 He always compares the maximum number of seeds produced by two species when crossed and by their hybrid offspring; with the average number produced by both pure parent…species in a state of nature。 But a serious cause of error seems to me to be here introduced: a plant to be hybridised must be castrated; and; what is often more important; must be secluded in order to prevent pollen being brought to it by insects from other plants。 Nearly all the plants experimentised on by G?rtner were potted; and apparently were kept in a chamber in his house。 That these processes are often injurious to the fertility of a plant cannot be doubted; for G?rtner gives in his table about a score of cases of plants which he castrated; and artificially fertilised with their own pollen; and (excluding all cases such as the Leguminosae; in which there is an acknowledged difficulty in the manipulation) half of these twenty plants had their fertility in some degree impaired。 Moreover; as G?rtner during several years repeatedly crossed the primrose and cowslip; which we have such good reason to believe to be varieties; and only once or twice succeeded in getting fertile seed; as he found the common red and blue pimpernels (Anagallis arvensis and coerulea); which the best botanists rank as varieties; absolutely sterile together; and as he came to the same conclusion in several other analogous cases; it seems to me that we may well be permitted to doubt whether many other species are really so sterile; when intercrossed; as G?rtner believes。 
It is certain; on the one hand; that the sterility of various species when crossed is so different in degree and graduates away so insensibly; and; on the other hand; that the fertility of pure species is so easily affected by various circumstances; that for all practical purposes it is most difficult to say where perfect fertility ends and sterility begins。 I think no better evidence of this can be required than that the two most experienced observers who have ever lived; namely; K?lreuter and G?rtner; should have arrived at diametrically opposite conclusions in regard to the very same species。 It is also most instructive to compare but I have not space here to enter on details the evidence advanced by our best botanists on the question whether certain doubtful forms should be ranked as species or varieties; with the evidence from fertility adduced by different hybridisers; or by the same author; from experiments made during different years。 It can thus be shown that neither sterility nor fertility affords any clear distinction between species and varieties; but that the evidence from this source graduates away; and is doubtful in the same degree as is the evidence derived from other constitutional and structural differences。 
In regard to the sterility of hybrids in successive generations; though G?rtner was enabled to rear some hybrids; carefully guarding them from a cross with either pure parent; for six or seven; and in one case for ten generations; yet he asserts positively that their fertility never increased; but generally greatly decreased。 I do not doubt that this is usually the case; and that the fertility often suddenly decreases in the first few generations。 Nevertheless I believe that in all these experiments the fertility has been diminished by an independent cause; namely; from close interbreeding。 I have collected so large a body of facts; showing that close interbreeding lessens fertility; and; on the other hand; that an occasional cross with a distinct individual or varie

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