Chapter 56

Reproduction may be divided into two main classes, namely, sexual and asexual. The latter is effected in many ways--by the formation of buds of various kinds, and by fissiparous generation, that is by spontaneous or artificial division. It is notorious that some of the lower animals, when cut into many pieces, reproduce so many perfect individuals: Lyonnet cut a Nais or freshwater worm into nearly forty pieces, and these all reproduced perfect animals. (27/2. Quoted by Paget 'Lectures on Pathology' 1853 page 159.) It is probable that segmentation could be carried much further in some of the protozoa; and with some of the lowest plants each cell will reproduce the parent-form. Johannes Muller thought that there was an important distinction between gemmation and fission; for in the latter case the divided portion, however small, is more fully developed than a bud, which also is a younger formation; but most physiologists are now convinced that the two processes are essentially alike. (27/3. Dr. Lachmann also observes ('Annals and Mag. of Nat. History' 2nd series volume 19 1857 page 231) with respect to infusoria, that "fissation and gemmation pass into each other almost imperceptibly." Again, Mr. W.C. Minor ('Annals and Mag. of Nat. Hist.' 3rd series volume 11 page 328) shows that with Annelids the distinction that has been made between fission and budding is not a fundamental one. See also Professor Clark's work 'Mind in Nature' New York 1865 pages 62, 94.) Prof. Huxley remarks, "fission is little more than a peculiar mode of budding," and Prof. H.J. Clark shows in detail that there is sometimes "a compromise between self-division and budding." When a limb is amputated, or when the whole body is bisected, the cut extremities are said to bud forth (27/4. See Bonnet 'Oeuvres d'Hist. Nat.' tome 5 1781 page 339 for remarks on the budding-out of the amputated limbs of Salamanders.); and as the papilla, which is first formed, consists of undeveloped cellular tissue like that forming an ordinary bud, the expression is apparently correct. We see the connection of the two processes in another way; for Trembley observed with the hydra, that the reproduction of the head after amputation was checked as soon as the animal put forth reproductive gemmae. (27/5. Paget 'Lectures on Pathology' 1853 page 158.)

Between the production, by fissiparous generation, of two or more complete individuals, and the repair of even a very slight injury, there is so perfect a gradation, that it is impossible to doubt that the two processes are connected. As at each stage of growth an amputated part is replaced by one in the same state of development, we must also follow Sir J. Paget in admitting, "that the powers of development from the embryo, are identical with those exercised for the restoration from injuries: in other words, that the powers are the same by which perfection is first achieved, and by which, when lost, it is recovered." (27/6. Ibid pages 152, 164.) Finally, we may conclude that the several forms of budding, fissiparous generation, the repair of injuries, and development, are all essentially the results of one and the same power.

SEXUAL GENERATION.

The union of the two sexual elements seems at first sight to make a broad distinction between sexual and asexual generation. But the conjugation of algae, by which process the contents of two cells unite into a single mass capable of development, apparently gives us the first step towards sexual union: and Pringsheim, in his memoir on the pairing of Zoospores (27/7. Translated in 'Annals and Mag. of Nat. Hist.' April 1870 page 272.), shows that conjugation graduates into true sexual reproduction. Moreover, the now well-ascertained cases of Parthenogenesis prove that the distinction between sexual and asexual generation is not nearly so great as was formerly thought; for ova occasionally, and even in some cases frequently, become developed into perfect beings, without the concourse of the male. With most of the lower animals and even with mammals, the ova show a trace of parthenogenetic power, for without being fertilised they pass through the first stages of segmentation. (27/8. Bischoff as quoted by von Siebold "Ueber Parthenogenesis" 'Sitzung der math. phys. Classe.' Munich November 4, 1871 page 240. See also Quatrefages 'Annales des Sc. Nat. Zoolog.' 3rd series 1850 page 138.) Nor can pseudova which do not need fertilisation, be distinguished from true ova, as was first shown by Sir J. Lubbock, and is now admitted by Siebold. So, again, the germ-balls in the larvae of Cecidomyia are said by Leuckart (27/9. 'On the Asexual Reproduction of Cecidomyide Larvae' translated in 'Annals and Mag. of Nat. Hist.' March 1866 pages 167, 171.) to be formed within the ovarium, but they do not require to be fertilised. It should also be observed that in sexual generation, the ovules and the male element have equal power of transmitting every single character possessed by either parent to their offspring. We see this clearly when hybrids are paired inter se, for the characters of both grandparents often appear in the progeny, either perfectly or by segments. It is an error to suppose that the male transmits certain characters and the female other characters; although no doubt, from unknown causes, one sex sometimes has a much stronger power of transmission than the other.

It has, however, been maintained by some authors that a bud differs essentially from a fertilised germ, in always reproducing the perfect character of the parent-stock; whilst fertilised germs give birth to variable beings. But there is no such broad distinction as this. In the eleventh chapter numerous cases were advanced showing that buds occasionally grow into plants having quite new characters; and the varieties thus produced can be propagated for a length of time by buds, and occasionally by seed. Nevertheless, it must be admitted that beings produced sexually are much more liable to vary than those produced asexually; and of this fact a partial explanation will hereafter be attempted. The variability in both cases is determined by the same general causes, and is governed by the same laws. Hence new varieties arising from buds cannot be distinguished from those arising from seed. Although bud-varieties usually retain their character during successive bud-generations, yet they occasionally revert, even after a long series of bud-generations, to their former character. This tendency to reversion in buds, is one of the most remarkable of the several points of agreement between the offspring from bud and seminal reproduction.

But there is one difference between organisms produced sexually and asexually, which is very general. The former pass in the course of their development from a very low stage to their highest stage, as we see in the metamorphoses of insects and of many other animals, and in the concealed metamorphoses of the vertebrata. Animals propagated asexually by buds or fission, on the other hand, commence their development at that stage at which the budding or self- dividing animal may happen to be, and therefore do not pass through some of the lower developmental stages. (27/10. Prof. Allman speaks ('Transact. R. Soc. of Edinburgh' volume 26 1870 page 102) decisively on this head with respect to the Hydroida: he says, "It is a universal law in the succession of zooids, that no retrogression ever takes place in the series.") Afterwards, they often advance in organisation, as we see in the many cases of "alternate generation." In thus speaking of alternate generation, I follow those naturalists who look at this process as essentially one of internal budding or of fissiparous generation. Some of the lower plants, however, such as mosses and certain algae, according to Dr. L. Radlkofer (27/11. 'Annals and Mag. of Nat. Hist.' 2nd series volume 20 1857 pages 153-455), when propagated asexually, do undergo a retrogressive metamorphosis. As far as the final cause is concerned, we can to a certain extent understand why beings propagated by buds should not pass through all the early stages of development; for with each organism the structure acquired at each stage must be adapted to its peculiar habits; and if there are places for the support of many individuals at some one stage, the simplest plan will be that they should be multiplied at this stage, and not that they should first retrograde in their development to an earlier or simpler structure, which might not be fitted for the then surrounding conditions.

From the several foregoing considerations we may conclude that the difference between sexual and asexual generation is not nearly so great as at first appears; the chief difference being that an ovule cannot continue to live and to be fully developed unless it unites with the male element; but even this difference is far from invariable, as shown by the many cases of parthenogenesis. We are therefore naturally led to inquire what the final cause can be of the necessity in ordinary generation for the concourse of the two sexual elements.

Seeds and ova are often highly serviceable as the means of disseminating plants and animals, and of preserving them during one or more seasons in a dormant state; but unimpregnated seeds or ova, and detached buds, would be equally serviceable for both purposes. We can, however, indicate two important advantages gained by the concourse of the two sexes, or rather of two individuals belonging to opposite sexes; for, as I have shown in a former chapter, the structure of every organism appears to be especially adapted for the concurrence, at least occasionally, of two individuals. When species are rendered highly variable by changed conditions of life, the free intercrossing of the varying individuals tends to keep each form fitted for its proper place in nature; and crossing can be effected only by sexual generation; but whether the end thus gained is of sufficient importance to account for the first origin of sexual intercourse is extremely doubtful. Secondly, I have shown from a large body of facts, that, as a slight change in the conditions of life is beneficial to each creature, so, in an analogous manner, is the change effected in the germ by sexual union with a distinct individual; and I have been led, from observing the many widely-extended provisions throughout nature for this purpose, and from the greater vigour of crossed organisms of all kinds, as proved by direct experiments, as well as from the evil effects of close interbreeding when long continued, to believe that the advantage thus gained is very great.

Why the germ, which before impregnation undergoes a certain amount of development, ceases to progress and perishes, unless it be acted on by the male element; and why conversely the male element, which in the case of some insects is enabled to keep alive for four or five years, and in the case of some plants for several years, likewise perishes, unless it acts on or unites with the germ, are questions which cannot be answered with certainty. It is, however, probable that both sexual elements perish, unless brought into union, simply from including too little formative matter for independent development. Quatrefages has shown in the case of the Teredo (27/12. 'Annales des Sc. Nat.' 3rd series 1850 tome 13.), as did formerly Prevost and Dumas with other animals, that more than one spermatozoon is requisite to fertilise an ovum. This has likewise been shown by Newport (27/13. 'Transact. Phil. Soc.' 1851 pages 196, 208, 210; 1853 pages 245, 247.), who proved by numerous experiments, that, when a very small number of spermatozoa are applied to the ova of Batrachians, they are only partially impregnated, and an embryo is never fully developed. The rate also of the segmentation of the ovum is determined by the number of the spermatozoa. With respect to plants, nearly the same results were obtained by Kolreuter and Gartner. This last careful observer, after making successive trials on a Malva with more and more pollen- grains, found (27/14. 'Beitrage zur Kenntniss' etc. 1844 s. 345.), that even thirty grains did not fertilise a single seed; but when forty grains were applied to the stigma, a few seeds of small size were formed. In the case of Mirabilis the pollen grains are extraordinarily large, and the ovarium contains only a single ovule; and these circumstances led Naudin (27/15. 'Nouvelles Archives du Museum' tome 1 page 27.) to make the following experiments: a flower was fertilised by three grains and succeeded perfectly; twelve flowers were fertilised by two grains, and seventeen flowers by a single grain, and of these one flower alone in each lot perfected its seed: and it deserves especial notice that the plants produced by these two seeds never attained their proper dimensions, and bore flowers of remarkably small size. From these facts we clearly see that the quantity of the peculiar formative matter which is contained within the spermatozoa and pollen-grains is an all-important element in the act of fertilisation, not only for the full development of the seed, but for the vigour of the plant produced from such seed. We see something of the same kind in certain cases of parthenogenesis, that is, when the male element is wholly excluded; for M. Jourdan (27/16. As quoted by Sir J. Lubbock in 'Nat. Hist. Review' 1862 page 345. Weijenbergh also raised ('Nature' December 21, 1871 page 149) two successive generations from unimpregnated females of another lepidopterous insect, Liparis dispar. These females did not produce at most one-twentieth of their full complement of eggs, and many of the eggs were worthless. Moreover the caterpillars raised from these unfertilised eggs "possessed far less vitality" than those from fertilised eggs. In the third parthenogenetic generation not a single egg yielded a caterpillar.) found that, out of about 58,000 eggs laid by unimpregnated silk-moths, many passed through their early embryonic stages, showing that they were capable of self-development, but only twenty-nine out of the whole number produced caterpillars. The same principle of quantity seems to hold good even in artificial fissiparous reproduction, for Hackel (27/17. 'Entwickelungsgeschichte der Siphonophora' 1869 page 73.) found that by cutting the segmented and fertilised ova or larva of Siphonophorae (jelly- fishes) into pieces, the smaller the pieces were, the slower was the rate of development, and the larvae thus produced were by so much the more imperfect and inclined to monstrosity. It seems, therefore, probable that with the separate sexual elements deficient quantity of formative matter is the main cause of their not having the capacity for prolonged existence and development, unless they combine and thus increase each other's bulk. The belief that it is the function of the spermatozoa to communicate life to the ovule seems a strange one, seeing that the unimpregnated ovule is already alive and generally undergoes a certain amount of independent development. Sexual and asexual reproduction are thus seen not to differ essentially; and we have already shown that asexual reproduction, the power of regrowth and development are all parts of one and the same great law.

REGROWTH OF AMPUTATED PARTS.

This subject deserves a little further discussion. A multitude of the lower animals and some vertebrates possess this wonderful power. For instance, Spallanzani cut off the legs and tail of the same salamander six times successively, and Bonnet (27/18. Spallanzani 'An Essay on Animal Reproduction' translated by Dr. Maty 1769 page 79. Bonnet 'Oeuvres d'Hist. Nat.' tome 5 part 1 4to. edition 1781 pages 343, 350.) did so eight times; and on each occasion the limbs were reproduced on the exact line of amputation, with no part deficient or in excess. An allied animal, the axolotl, had a limb bitten off, which was reproduced in an abnormal condition, but when this was amputated it was replaced by a perfect limb. (27/19. Vulpian as quoted by Prof. Faivre 'La Variabilite des Especes' 1868 page 112.) The new limbs in these cases bud forth, and are developed in the same manner as during the regular development of a young animal. For instance, with the Amblystoma lurida, three toes are first developed, then the fourth, and on the hind-feet the fifth, and so it is with a reproduced limb. (27/20. Dr. P. Hoy 'The American Naturalist' September 1871 page 579.)

The power of regrowth is generally much greater during the youth of an animal or during the earlier stages of its development than during maturity. The larvae or tadpoles of the Batrachians are capable of reproducing lost members, but not so the adults. (27/21. Dr. Gunther in Owen 'Anatomy of Vertebrates'