According to the belief now generally accepted by our best naturalists, all the members of the same order or class, the Macrourous crustaceans for instance, are descended from a common progenitor. During their descent they have diverged much in structure, but have retained much in common; and this divergence and retention of character has been effected, though they have passed and still pass through marvellously different metamorphoses. This fact well illustrates how independent each structure must be from that which precedes and follows it in the course of development.
The Functional Independence of the Elements or Units of the Body.—Physiologists agree that the whole organism consists of a multitude of elemental parts, which are to a great extent independent of each other. Each organ, says Claude Bernard,[891]has its proper life, its autonomy; it can develop and reproduce itself independently of the adjoining tissues. The great German authority, Virchow,[892]asserts still more emphatically that each system, as the nervous or osseous system, or the blood, consists of an "enormous mass of minute centres of action.... Every element has its own special action, and even though it derive its stimulus to activity from other parts, yet alone effects the actual performance of its duties.... Every single epithelial and muscular fibre-cell leads a sort of parasitical existence in relation to the rest of the body.... Every single bone-corpuscle really possesses conditions of nutrition peculiar to itself." Each element, as Mr. Paget remarks, lives its appointed time, and then dies, and, after being cast off or absorbed, is replaced.[893]I presume that no physiologist doubts that, for instance, each bone-corpuscle of the finger differs from the corresponding corpuscle in the corresponding joint of the toe; and there can hardly be a doubt that even those on the corresponding sides of the body differ, though almost identical in nature. This near approach to identity is curiously shown in many diseases in which the same exact points on the right and left sides of the body are similarly affected; thus Mr. Paget[894]gives a drawing of a diseased pelvis, in which the bone has grown into a most complicated pattern, but "there is not one spot or line on one side which is not represented, as exactly as it would be in a mirror, on the other."
Many facts support this view of the independent life of each minute element of the body. Virchow insists that a single bone-corpuscle or a single cell in the skin may become diseased. The spur of a cock, after being inserted into the eye of an ox, lived for eight years, and acquired a weight of 306 grammes, or nearly fourteen ounces.[895]The tail of a pig has been grafted into the middle of its back, and reacquired sensibility. Dr. Ollier[896]inserted a piece of periosteum from the bone of a young dog under the skin of a rabbit, and true bone was developed. A multitude of similar facts could be given. Thefrequent presence of hairs and of perfectly developed teeth, even teeth of the second dentition, in ovarian tumours,[897]are facts leading to the same conclusion.
Whether each of the innumerable autonomous elements of the body is a cell or the modified product of a cell, is a more doubtful question, even if so wide a definition be given to the term, as to include cell-like bodies without walls and without nuclei.[898]Professor Lionel Beale uses the term "germinal matter" for the contents of cells, taken in this wide acceptation, and he draws a broad distinction between germinal matter and "formed material" or the various products of cells.[899]But the doctrine ofomnis cellula e cellulâis admitted for plants, and is a widely prevalent belief with respect to animals.[900]Thus Virchow, the great supporter of the cellular theory, whilst allowing that difficulties exist, maintains that every atom of tissue is derived from cells, and these from pre-existing cells, and these primarily from the egg, which he regards as a great cell. That cells, still retaining the same nature, increase by self-division or proliferation, is admitted by almost every one. But when an organism undergoes a great change of structure during development, the cells, which at each stage are supposed to be directly derived from previously-existing cells, must likewise be greatly changed in nature; this change is apparently attributed by the supporters of the cellular doctrine to some inherent power which the cells possess, and not to any external agency.
Another school maintains that cells and tissues of all kinds may be formed, independently of pre-existing cells, from plastic lymph or blastema; and this it is thought is well exhibited in the repair of wounds. As I have not especially attended to histology, it would be presumptuous in me to express an opinion on the two opposed doctrines. But every one appears to admit that the body consists of a multitude of "organic units,"[901]each of which possesses its own proper attributes, and is to a certain extent independent of all others. Hence it will be convenient to use indifferently the terms cells or organic units or simply units.
Variability and Inheritance.—We have seen in the twenty-second chapter that variability is not a principle co-ordinate with life or reproduction, but results from special causes, generally from changed conditions acting during successive generations. Part of the fluctuating variability thus induced is apparently due to the sexual system being easily affected by changed conditions, so that it is often rendered impotent; and when not so seriously affected, it often fails in its proper function of transmitting truly the characters of the parents to the offspring. But variability is not necessarily connected with the sexual system, as we see from the cases of bud-variation; and although we may not be able to trace the nature of the connexion, it is probable that many deviations of structure which appear in sexual offspring result from changed conditions acting directly on the organisation, independently of the reproductive organs. In some instances we may feel sure of this, when all, or nearly all the individuals which have been similarly exposed are similarly and definitely affected—as in the dwarfed and otherwise changed maize brought from hot countries when cultivated in Germany; in the change of the fleece in sheep within the tropics; to a certain extent in the increased size and early maturity of our highly-improved domesticated animals; in inherited gout from intemperance; and in many other such cases. Now, as such changed conditions do not especially affect the reproductive organs, it seems mysterious on any ordinary view why their product, the new organic being, should be similarly affected.
How, again, can we explain to ourselves the inherited effects of the use or disuse of particular organs? The domesticated duck flies less and walks more than the wild duck, and its limb-bones have become in a corresponding manner diminished and increased in comparison with those of the wild duck. A horse is trained to certain paces, and the colt inherits similar consensual movements. The domesticated rabbit becomes tame from close confinement; the dog intelligent from associating with man; the retriever is taught to fetch and carry: and thesemental endowments and bodily powers are all inherited. Nothing in the whole circuit of physiology is more wonderful. How can the use or disuse of a particular limb or of the brain affect a small aggregate of reproductive cells, seated in a distant part of the body, in such a manner that the being developed from these cells inherits the characters of either one or both parents? Even an imperfect answer to this question would be satisfactory.
Sexual reproduction does not essentially differ, as we have seen, from budding or self-division, and these processes graduate through the repair of injuries into ordinary development and growth; it might therefore be expected that every character would be as regularly transmitted by all the methods of reproduction as by continued growth. In the chapters devoted to inheritance it was shown that a multitude of newly-acquired characters, whether injurious or beneficial, whether of the lowest or highest vital importance, are often faithfully transmitted—frequently even when one parent alone possesses some new peculiarity. It deserves especial attention that characters appearing at any age tend to reappear at a corresponding age. We may on the whole conclude that in all cases inheritance is the rule, and non-inheritance the anomaly. In some instances a character is not inherited, from the conditions of life being directly opposed to its development; in many instances, from the conditions incessantly inducing fresh variability, as with grafted fruit-trees and highly cultivated flowers. In the remaining cases the failure may be attributed to reversion, by which the child resembles its grandparents or more remote progenitors, instead of its parents.
This principle of Reversion is the most wonderful of all the attributes of Inheritance. It proves to us that the transmission of a character and its development, which ordinarily go together and thus escape discrimination, are distinct powers; and these powers in some cases are even antagonistic, for each acts alternately in successive generations. Reversion is not a rare event, depending on some unusual or favourable combination of circumstances, but occurs so regularly with crossed animals and plants, and so frequently with uncrossed breeds, that it is evidently an essential part of the principle of inheritance. We know thatchanged conditions have the power of evoking long-lost characters, as in the case of some feral animals. The act of crossing in itself possesses this power in a high degree. What can be more wonderful than that characters, which have disappeared during scores, or hundreds, or even thousands of generations, should suddenly reappear perfectly developed, as in the case of pigeons and fowls when purely bred, and especially when crossed; or as with the zebrine stripes on dun-coloured horses, and other such cases? Many monstrosities come under this same head, as when rudimentary organs are redeveloped, or when an organ which we must believe was possessed by an early progenitor, but of which not even a rudiment is left, suddenly reappears, as with the fifth stamen in some Scrophulariaceæ. We have already seen that reversion acts in bud-reproduction; and we know that it occasionally acts during the growth of the same individual animal, especially, but not exclusively, when of crossed parentage,—as in the rare cases described of individual fowls, pigeons, cattle, and rabbits, which have reverted as they advanced in years to the colours of one of their parents or ancestors.
We are led to believe, as formerly explained, that every character which occasionally reappears is present in a latent form in each generation, in nearly the same manner as in male and female animals secondary characters of the opposite sex lie latent, ready to be evolved when the reproductive organs are injured. This comparison of the secondary sexual characters which are latent in both sexes, with other latent characters, is the more appropriate from the case recorded of the Hen, which assumed some of the masculine characters, not of her own race, but of an early progenitor; she thus exhibited at the same time the redevelopment of latent characters of both kinds and connected both classes. In every living creature we may feel assured that a host of lost characters lie ready to be evolved under proper conditions. How can we make intelligible, and connect with other facts, this wonderful and common capacity of reversion,—this power of calling back to life long-lost characters?
Part II.
I have now enumerated the chief facts which every one would desire to connect by some intelligible bond. This can be done, as it seems to me, if we make the following assumptions; if the first and chief one be not rejected, the others, from being supported by various physiological considerations, will not appear very improbable. It is almost universally admitted that cells, or the units of the body, propagate themselves by self-division or proliferation, retaining the same nature, and ultimately becoming converted into the various tissues and substances of the body. But besides this means of increase I assume that cells, before their conversion into completely passive or "formed material," throw off minute granules or atoms, which circulate freely throughout the system, and when supplied with proper nutriment multiply by self-division, subsequently becoming developed into cells like those from which they were derived. These granules for the sake of distinctness may be called cell-gemmules, or, as the cellular theory is not fully established, simply gemmules. They are supposed to be transmitted from the parents to the offspring, and are generally developed in the generation which immediately succeeds, but are often transmitted in a dormant state during many generations and are then developed. Their development is supposed to depend on their union with other partially developed cells or gemmules which precede them in the regular course of growth. Why I use the term union, will be seen when we discuss the direct action of pollen on the tissues of the mother-plant. Gemmules are supposed to be thrown off by every cell or unit, not only during the adult state, but during all the stages of development. Lastly, I assume that the gemmules in their dormant state have a mutual affinity for each other, leading to their aggregation either into buds or into the sexual elements. Hence, speaking strictly, it is not the reproductive elements, nor the buds, which generate new organisms, but the cells themselves throughout the body. These assumptions constitute the provisional hypothesis which I have called Pangenesis. Nearlysimilar views have been propounded, as I find, by other authors, more especially by Mr. Herbert Spencer;[902]but they are here modified and amplified.
Before proceeding to show, firstly, how far these assumptions are in themselves probable, and secondly, how far they connect and explain the various groups of facts with which we are concerned, it may be useful to give an illustration of the hypothesis. If one of the simplest Protozoa be formed, as appears under the microscope, of a small mass of homogeneous gelatinous matter, a minute atom thrown off from any part and nourished under favourable circumstances would naturally reproduce the whole; but if the upper and lower surfaces were to differ in texture from the central portion, then all three parts would have to throw off atoms or gemmules, which when aggregated by mutual affinity would form either buds or the sexual elements. Precisely the same view may be extended to one of the higher animals; although in this case many thousand gemmules must be thrown off from the various parts of the body. Now, when the leg, for instance, of a salamander is cut off, a slight crust forms over the wound, and beneath this crust the uninjured cells or units of bone, muscle, nerves, &c., are supposed to unite with the diffused gemmules of those cells which in the perfect leg come next in order; and these as they become slightly developed unite with others, and so on until a papilla of soft cellular tissue, the "budding leg," is formed, and in time a perfect leg.[903]Thus, that portion of the leg which hadbeen cut off, neither more nor less, would be reproduced. If the tail or leg of a young animal had been cut off, a young tail or leg would have been reproduced, as actually occurs with the amputated tail of the tadpole; for gemmules of all the units which compose the tail are diffused throughout the body at all ages. But during the adult state the gemmules of the larval tail would remain dormant, for they would not meet with pre-existing cells in a proper state of development with which to unite. If from changed conditions or any other cause any part of the body should become permanently modified, the gemmules, which are merely minute portions of the contents of the cells forming the part, would naturally reproduce the same modification. But gemmules previously derived from the same part before it had undergone any change, would still be diffused throughout the organisation, and would be transmitted from generation to generation, so that under favourable circumstances they might be redeveloped, and then the new modification would be for a time or for ever lost. The aggregation of gemmules derived from every part of the body, through their mutual affinity, would form buds, and their aggregation in some special manner, apparently in small quantity, together probably with the presence of gemmules of certain primordial cells, would constitute the sexual elements. By means of these illustrations the hypothesis of pangenesis has, I hope, been rendered intelligible.
Physiologists maintain, as we have seen, that each cell, though to a large extent dependent on others, is likewise, to a certain extent, independent or autonomous. I go one small step further, and assume that each cell casts off a free gemmule, which is capable of reproducing a similar cell. There is some analogy between this view and what we see in compound animals and in the flower-buds on the same tree; for these are distinct individuals capable of true or seminal reproduction, yet have parts in common and are dependent on each other; thusthe tree has its bark and trunk, and certain corals, as the Virgularia, have not only parts, but movements in common.
The existence of free gemmules is a gratuitous assumption, yet can hardly be considered as very improbable, seeing that cells have the power of multiplication through the self-division of their contents. Gemmules differ from true ovules or buds inasmuch as they are supposed to be capable of multiplication in their undeveloped state. No one probably will object to this capacity as improbable. The blastema within the egg has been known to divide and give birth to two embryos; and Thuret[904]has seen the zoospore of an alga divide itself, and both halves germinate. An atom of small-pox matter, so minute as to be borne by the wind, must multiply itself many thousand-fold in a person thus inoculated.[905]It has recently been ascertained[906]that a minute portion of the mucous discharge from an animal affected with rinderpest, if placed in the blood of a healthy ox, increases so fast that in a short space of time "the whole mass of blood, weighing many pounds, is infected, and every small particle of that blood contains enough poison to give, within less than forty-eight hours, the disease to another animal."
The retention of free and undeveloped gemmules in the same body from early youth to old age may appear improbable, but we should remember how long seeds lie dormant in the earth and buds in the bark of a tree. Their transmission from generation to generation may appear still more improbable; but here again we should remember that many rudimentary and useless organs are transmitted and have been transmitted during an indefinite number of generations. We shall presently see how well the long-continued transmission of undeveloped gemmules explains many facts.
As each unit, or group of similar units throughout the body, casts off its gemmules, and as all are contained within the smallest egg or seed, and within each spermatozoon or pollen-grain, their number and minuteness must be somethinginconceivable. I shall hereafter recur to this objection, which at first appears so formidable; but it may here be remarked that a cod-fish has been found to produce 4,872,000 eggs, a single Ascaris about 64,000,000 eggs, and a single Orchidaceous plant probably as many million seeds.[907]In these several cases, the spermatozoa and pollen-grains must exist in considerably larger numbers. Now, when we have to deal with numbers such as these, which the human intellect cannot grasp, there is no good reason for rejecting our present hypothesis on account of the assumed existence of cell-gemmules a few thousand times more numerous.
The gemmules in each organism must be thoroughly diffused; nor does this seem improbable considering their minuteness, and the steady circulation of fluids throughout the body. So it must be with the gemmules of plants, for with certain kinds even a minute fragment of a leaf will reproduce the whole. But a difficulty here occurs; it would appear that with plants, and probably with compound animals, such as corals, the gemmules do not spread from bud to bud, but only through the tissues developed from each separate bud. We are led to this conclusion from the stock being rarely affected by the insertion of a bud or graft from a distinct variety. This non-diffusion of the gemmules is still more plainly shown in the case of ferns; for Mr. Bridgman[908]has proved that, when spores (which it should be remembered are of the nature of buds) are taken from a monstrous part of a frond, and others from an ordinary part,each reproduces the form of the part whence derived. But this non-diffusion of the gemmules from bud to bud may be only apparent, depending, as we shall hereafter see, on the nature of the first-formed cells in the buds.
The assumed elective affinity of each gemmule for that particular cell which precedes it in the order of development is supported by many analogies. In all ordinary cases of sexual reproduction the male and female elements have a mutual affinity for each other: thus, it is believed that about ten thousand species of Compositæ exist, and there can be no doubt that if the pollen of all these species could be, simultaneously or successively, placed on the stigma of any one species, this one would elect with unerring certainty its own pollen. This elective capacity is all the more wonderful, as it must have been acquired since the many species of this great group of plants branched off from a common progenitor. On any view of the nature of sexual reproduction, the protoplasm contained within the ovules and within the sperm-cells (or the "spermatic force" of the latter, if so vague a term be preferred) must act on each other by some law of special affinity, either during or subsequently to impregnation, so that corresponding parts alone affect each other; thus, a calf produced from a short-horned cow by a long-horned bull has its horns and not its horny hoofs affected by the union of the two forms, and the offspring from two birds with differently coloured tails have their tails and not their whole plumage affected.
The various tissues of the body plainly show, as many physiologists have insisted,[909]an affinity for special organic substances, whether natural or foreign to the body. We see this in the cells of the kidneys attracting urea from the blood; in the worrara poison affecting the nerves; upas and digitalis the muscles; the Lytta vesicatoria the kidneys; and in the poisonous matter of many diseases, as small-pox, scarlet-fever, hooping-cough, glanders, cancer, and hydrophobia, affecting certain definite parts of the body or certain tissues or glands.
The affinity of various parts of the body for each other duringtheir early development was shown in the last chapter, when discussing the tendency to fusion in homologous parts. This affinity displays itself in the normal fusion of organs which are separate at an early embryonic age, and still more plainly in those marvellous cases of double monsters in which each bone, muscle, vessel, and nerve in the one embryo, blends with the corresponding part in the other. The affinity between homologous organs may come into action with single parts, or with the entire individual, as in the case of flowers or fruits which are symmetrically blended together with all their parts doubled, but without any other trace of fusion.
It has also been assumed that the development of each gemmule depends on its union with another cell or unit which has just commenced its development, and which, from preceding it in order of growth, is of a somewhat different nature. Nor is it a very improbable assumption that the development of a gemmule is determined by its union with a cell slightly different in nature, for abundant evidence was given in the seventeenth chapter, showing that a slight degree of differentiation in the male and female sexual elements favours in a marked manner their union and subsequent development. But what determines the development of the gemmules of the first-formed or primordial cell in the unimpregnated ovule, is beyond conjecture.
It must also be admitted that analogy fails to guide us towards any determination on several other points: for instance, whether cells, derived from the same parent-cell, may, in the regular course of growth, become developed into different structures, from absorbing peculiar kinds of nutriment, independently of their union with distinct gemmules. We shall appreciate this difficulty if we call to mind, what complex yet symmetrical growths the cells of plants yield when they are inoculated by the poison of a gall-insect. With animals various polypoid excrescences and tumours are now generally admitted[910]to be the direct product, through proliferation, of normal cells which have become abnormal. In the regular growth and repair of bones, the tissues undergo, as Virchow remarks,[911]a whole series of permutations and substitutions. "The cartilage-cells may beconverted by a direct transformation into marrow-cells, and continue as such; or they may first be converted into osseous and then into medullary tissue; or lastly, they may first be converted into marrow and then into bone. So variable are the permutations of these tissues, in themselves so nearly allied, and yet in their external appearance so completely distinct." But as these tissues thus change their nature at any age, without any obvious change in their nutrition, we must suppose in accordance with our hypothesis that gemmules derived from one kind of tissue combine with the cells of another kind, and cause the successive modifications.
It is useless to speculate at what period of development each organic unit casts off its gemmules; for the whole subject of the development of the various elemental tissues is as yet involved in much doubt. Some physiologists, for instance, maintain that muscle or nerve-fibres are developed from cells, which are afterwards nourished by their own proper powers of absorption; whilst other physiologists deny their cellular origin; and Beale maintains that such fibres are renovated exclusively by the conversion of fresh germinal matter (that is the so-called nuclei) into "formed material." However this may be, it appears probable that all external agencies, such as changed nutrition, increased use or disuse, &c., which induced any permanent modification in a structure, would at the same time or previously act on the cells, nuclei, germinal or formative matter, from which the structures in question were developed, and consequently would act on the gemmules or cast-off atoms.
There is another point on which it is useless to speculate, namely, whether all gemmules are free and separate, or whether some are from the first united into small aggregates. A feather, for instance, is a complex structure, and, as each separate part is liable to inherited variations, I conclude that each feather certainly generates a large number of gemmules; but it is possible that these may be aggregated into a compound gemmule. The same remark applies to the petals of a flower, which in some cases are highly complex, with each ridge and hollow contrived for special purposes, so that each part must have been separately modified, and the modifications transmitted; consequently, separate gemmules, according to our hypothesis,must have been thrown off from each cell or part. But, as we sometimes see half an anther or a small portion of a filament becoming petaliform, or parts or mere stripes of the calyx assuming the colour and texture of the corolla, it is probable that with petals the gemmules of each cell are not aggregated together into a compound gemmule, but are freely and separately diffused.
Having now endeavoured to show that the several foregoing assumptions are to a certain extent supported by analogous facts, and having discussed some of the most doubtful points, we will consider how far the hypothesis brings under a single point of view the various cases enumerated in the First Part. All the forms of reproduction graduate into each other and agree in their product; for it is impossible to distinguish between organisms produced from buds, from self-division, or from fertilised germs; such organisms are liable to variations of the same nature and to reversion of character; and as we now see that all the forms of reproduction depend on the aggregation of gemmules derived from the whole body, we can understand this general agreement. It is satisfactory to find that sexual and asexual generation, by both of which widely different processes the same living creature is habitually produced, are fundamentally the same. Parthenogenesis is no longer wonderful; in fact, the wonder is that it should not oftener occur. We see that the reproductive organs do not actually create the sexual elements; they merely determine or permit the aggregation of the gemmules in a special manner. These organs, together with their accessory parts, have, however, high functions to perform; they give to both elements a special affinity for each other, independently of the contents of the male and female cells, as is shown in the case of plants by the mutual reaction of the stigma and pollen-grains; they adapt one or both elements for independent temporary existence, and for mutual union. The contrivances for these purposes are sometimes wonderfully complex, as with the spermatophores of the Cephalopoda. The male element sometimes possesses attributes which, if observed in an independent animal, would be put down to instinct guided by sense-organs, as when thespermatozoon of an insect finds its way into the minute micropyle of the egg, or as when the antherozoids of certain algæ swim by the aid of their ciliæ to the female plant, and force themselvesintoa minute orifice. In these latter cases, however, we must believe that the male element has acquired its powers, on the same principle with the larvæ of animals, namely by successive modifications developed at corresponding periods of life: we can hardly avoid in these cases looking at the male element as a sort of premature larva, which unites, or, like one of the lower algæ, conjugates, with the female element. What determines the aggregation of the gemmules within the sexual organs we do not in the least know; nor do we know why buds are formed in certain definite places, leading to the symmetrical growth of trees and corals, nor why adventitious buds may be formed almost anywhere, even on a petal, and frequently upon healed wounds.[912]As soon as the gemmules have aggregated themselves, development apparently commences, but in the case of buds is often afterwards suspended, and in the case of the sexual elements soon ceases, unless the elements of the opposite sexes combine; even after this has occurred, the fertilised germ, as with seeds buried in the ground, may remain during a lengthened period in a dormant state.
The antagonism which has long been observed,[913]though exceptions occur,[914]between active growth and the power of sexual reproduction—between the repair of injuries and gemmation—and with plants, between rapid increase by buds, rhizomes, &c., and the production of seed, is partly explained by the gemmules not existing in sufficient numbers for both processes.But this explanation hardly applies to those plants which naturally produce a multitude of seeds, but which, through a comparatively small increase in the number of the buds on their rhizomes or offsets, yield few or no seed. As, however, we shall presently see that buds probably include tissue which has already been to a certain extent developed or differentiated, some additional organised matter will thus have been expended.
From one of the forms of Reproduction, namely, spontaneous self-division, we are led by insensible steps to the repair of the slightest injury; and the existence of gemmules, derived from every cell or unit throughout the body and everywhere diffused, explains all such cases,—even the wonderful fact that, when the limbs of the salamander were cut off many times successively by Spallanzani and Bonnet, they were exactly and completely reproduced. I have heard this process compared with the recrystallisation which occurs when the angles of a broken crystal are repaired; and the two processes have this much in common, that in the one case the polarity of the molecules is the efficient cause, and in the other the affinity of the gemmules for particular nascent cells.
Pangenesis does not throw much light on Hybridism, but agrees well with most of the ascertained facts. We may conclude from the fact of a single spermatozoon or pollen-grain being insufficient for impregnation, that a certain number of gemmules derived from each cell or unit are required for the development of each part. From the occurrence of parthenogenesis, more especially in the case of the silk-moth, in which the embryo is often partially formed, we may also infer that the female element includes nearly sufficient gemmules of all kinds for independent development, so that when united with the male element the gemmules must be superabundant. Now, as a general rule, when two species or races are crossed reciprocally, the offspring do not differ, and this shows that both sexual elements agree in power, in accordance with the view that they include the same gemmules. Hybrids and mongrels are generally intermediate in character between the two parent-forms, yet occasionally they closely resemble one parent in one part and the other parent in another part, or even in their whole structure: nor is this difficult to understand onthe admission that the gemmules in the fertilised germ are superabundant in number, and that those derived from one parent have some advantage in number, affinity, or vigour over those derived from the other parent. Crossed forms sometimes exhibit the colour or other characters of either parent in stripes or blotches; and this may occur in the first generation, or through reversion in succeeding bud and seminal generations, as in the several instances given in the eleventh chapter. In these cases we must follow Naudin,[915]and admit that the "essence" or "element" of the two species, which terms I should translate into the gemmules, have an affinity for their own kind, and thus separate themselves into distinct stripes or blotches; and reasons were given, when discussing in the fifteenth chapter the incompatibility of certain characters to unite, for believing in such mutual affinity. When two forms are crossed, one is not rarely found to be prepotent in the transmission of character over the other; and this we can explain only by again assuming that the one form has some advantage in the number, vigour, or affinity of its gemmules, except in those cases, where certain characters are present in the one form and latent in the other. For instance, there is a latent tendency in all pigeons to become blue, and, when a blue pigeon is crossed with one of any other colour, the blue tint is generally prepotent. When we consider latent characters, the explanation of this form of prepotency will be obvious.
When one species is crossed with another it is notorious that they do not yield the full or proper number of offspring; and we can only say on this head that, as the development of each organism depends on such nicely-balanced affinities between a host of gemmules and developing cells or units, we need not feel at all surprised that the commixture of gemmules derived from two distinct species should lead to a partial or complete failure of development. With respect to the sterility of hybrids produced from the union of two distinct species, it was shown in the nineteenth chapter that this depends exclusively on the reproductive organs being specially affected; but why these organs should be thus affected we do not know, any more thanwhy unnatural conditions of life, though compatible with health, should cause sterility; or why continued close interbreeding, or the illegitimate unions of dimorphic and trimorphic plants, induce the same result. The conclusion that the reproductive organs alone are affected, and not the whole organisation, agrees perfectly with the unimpaired or even increased capacity in hybrid plants for propagation by buds; for this implies, according to our hypothesis, that the cells of the hybrids throw off hybridised cell-gemmules, which become aggregated into buds, but fail to become aggregated within the reproductive organs, so as to form the sexual elements. In a similar manner many plants, when placed under unnatural conditions, fail to produce seed, but can readily be propagated by buds. We shall presently see that pangenesis agrees well with the strong tendency to reversion exhibited by all crossed animals and plants.
It was shown in the discussion on graft-hybrids that there is some reason to believe that portions of cellular tissue taken from distinct plants become so intimately united, as afterwards occasionally to produce crossed or hybridised buds. If this fact were fully established, it would, by the aid of our hypothesis, connect gemmation and sexual reproduction in the closest manner.
Abundant evidence has been advanced proving that pollen taken from one species or variety and applied to the stigma of another sometimes directly affects the tissues of the mother-plant. It is probable that this occurs with many plants during fertilisation, but can only be detected when distinct forms are crossed. On any ordinary theory of reproduction this is a most anomalous circumstance, for the pollen-grains are manifestly adapted to act on the ovule, but in these cases they act on the colour, texture, and form of the coats of the seeds, on the ovarium itself, which is a modified leaf, and even on the calyx and upper part of the flower-peduncle. In accordance with the hypothesis of pangenesis pollen includes gemmules, derived from every part of the organisation, which diffuse themselves and multiply by self-division; hence it is not surprising that gemmules within the pollen, which are derived from the parts near the reproductive organs, should sometimes be able to affect the same parts, whilst still undergoing development, in the mother-plant.
As, during all the stages of development, the tissues of plants consist of cells, and as new cells are not known to be formed between, or independently of, pre-existing cells, we must conclude that the gemmules derived from the foreign pollen do not become developed merely in contact with pre-existing cells, but actually penetrate the nascent cells of the mother-plant. This process may be compared with the ordinary act of fertilisation, during which the contents of the pollen-tubes penetrate the closed embryonic sack within the ovule, and determine the development of the embryo. According to this view, the cells of the mother-plant may almost literally be said to be fertilised by the gemmules derived from the foreign pollen. With all organisms, as we shall presently see, the cells or organic units of the embryo during the successive stages of development may in like manner be said to be fertilised by the gemmules of the cells, which come next in the order of formation.
Animals, when capable of sexual reproduction, are fully developed, and it is scarcely possible that the male element should affect the tissues of the mother in the same direct manner as with plants; nevertheless it is certain that her ovaria are sometimes affected by a previous impregnation, so that the ovules subsequently fertilised by a distinct male are plainly influenced in character; and this, as in the case of foreign pollen, is intelligible through the diffusion, retention, and action of the gemmules included within the spermatozoa of the previous male.
Each organism reaches maturity through a longer or shorter course of development. The changes may be small and insensibly slow, as when a child grows into a man, or many, abrupt, and slight, as in the metamorphoses of certain ephemerous insects, or again few and strongly marked, as with most other insects. Each part may be moulded within a previously existing and corresponding part, and in this case it will appear, falsely as I believe, to be formed from the old part; or it may be developed within a wholly distinct part of the body, as in the extreme cases of metagenesis. An eye, for instance, may be developed at a spot where no eye previously existed. We have also seen that allied organic beings in the course of their metamorphoses sometimes attain nearly the same structure after passingthrough widely different forms; or conversely, after passing through nearly the same early forms, arrive at a widely different termination. In these cases it is very difficult to believe that the early cells or units possess the inherent power, independently of any external agent, of producing new structures wholly different in form, position, and function. But these cases become plain on the hypothesis of pangenesis. The organic units, during each stage of development, throw off gemmules, which, multiplying, are transmitted to the offspring. In the offspring, as soon as any particular cell or unit in the proper order of development becomes partially developed, it unites with (or to speak metaphorically is fertilised by) the gemmule of the next succeeding cell, and so onwards. Now, supposing that at any stage of development, certain cells or aggregates of cells had been slightly modified by the action of some disturbing cause, the cast-off gemmules or atoms of the cell-contents could hardly fail to be similarly affected, and consequently would reproduce the same modification. This process might be repeated until the structure of the part at this particular stage of development became greatly changed, but this would not necessarily affect other parts whether previously or subsequently developed. In this manner we can understand the remarkable independence of structure in the successive metamorphoses, and especially in the successive metageneses of many animals.
The term growth ought strictly to be confined to mere increase of size, and development to change of structure.[916]Now, a child is said to grow into a man, and a foal into a horse, but, as in these cases there is much change of structure, the process properly belongs to the order of development. We have indirect evidence of this in many variations and diseases supervening during so-called growth at a particular period, and being inherited at a corresponding period. In the case, however, of diseases which supervene during old age, subsequently to the ordinary period of procreation, and which nevertheless are sometimes inherited, as occurs with brain and heart complaints, wemust suppose that the organs were in fact affected at an earlier age and threw off at this period affected gemmules; but that the affection became visible or injurious only after the prolonged growth of the part in the strict sense of the word. In all the changes of structure which regularly supervene during old age, we see the effects of deteriorated growth, and not of true development.
In the so-called process ofalternate generationmany individuals are generated asexually during very early or later stages of development. These individuals may closely resemble the preceding larval form, but generally are wonderfully dissimilar. To understand this process we must suppose that at a certain stage of development the gemmules are multiplied at an unusual rate, and become aggregated by mutual affinity at many centres of attraction, or buds. These buds, it may be remarked, must include gemmules not only of all the succeeding but likewise of all the preceding stages of development; for when mature they have the power of transmitting by sexual generation gemmules of all the stages, however numerous these may be. It was shown in the First Part, at least in regard to animals, that the new beings which are thus at any period asexually generated do not retrograde in development—that is, they do not pass through those earlier stages, through which the fertilised germ of the same animal has to pass; and an explanation of this fact was attempted as far as the final or teleological cause is concerned. We can likewise understand the proximate cause, if we assume, and the assumption is far from improbable, that buds, like chopped-up fragments of a hydra, are formed of tissue which has already passed through several of the earlier stages of development; for in this case their component cells or units would not unite with the gemmules derived from the earlier-formed cells, but only with those which came next in the order of development. On the other hand, we must believe that, in the sexual elements, or probably in the female alone, gemmules of certain primordial cells are present; and these, as soon as their development commences, unite in due succession with the gemmules of every part of the body, from the first to the last period of life.
The principle of the independent formation of each part, inso far as its development depends on the union of the proper gemmules with certain nascent cells, together with the superabundance of the gemmules derived from both parents and self-multiplied, throws light on a widely different group of facts, which on any ordinary view of development appears very strange. I allude to organs which are abnormally multiplied or transposed. Thus gold-fish often have supernumerary fins placed on various parts of their bodies. We have seen that, when the tail of a lizard is broken off, a double tail is sometimes reproduced, and when the foot of the salamander is divided longitudinally, additional digits are occasionally formed. When frogs, toads, &c., are born with their limbs doubled, as sometimes occurs, the doubling, as Gervais remarks,[917]cannot be due to the complete fusion of two embryos, with the exception of the limbs, for the larvæ are limbless. The same argument is applicable[918]to certain insects produced with multiple legs or antennæ, for these are metamorphosed from apodal or antennæless larvæ. Alphonse Milne-Edwards[919]has described the curious case of a crustacean in which one eye-peduncle supported, instead of a complete eye, only an imperfect cornea, out of the centre of which a portion of an antenna was developed. A case has been recorded[920]of a man who had during both dentitions a double tooth in place of the left second incisor, and he inherited this peculiarity from his paternal grandfather. Several cases are known[921]of additional teeth having been developed in the palate, more especially with horses, and in the orbit of the eye. Certain breeds of sheep bear a whole crowd of horns on their foreheads. Hairs occasionally appear in strange situations, as within the ears of the Siamese hairy family; and hairs "quite natural in structure" have been observed "within the substance of the brain."[922]As many as five spurs have been seen on both legs in certain Game-fowls. In the Polish fowl the male is ornamented with a topknot of hackleslike those on his neck, whilst the female has one of common feathers. In feather-footed pigeons and fowls, feathers like those on the wing arise from the outer side of the legs and toes. Even the elemental parts of the same feather may be transposed; for in the Sebastopol goose, barbules are developed on the divided filaments of the shaft.
Analogous cases are of such frequent occurrence with plants that they do not strike us with sufficient surprise. Supernumerary petals, stamens, and pistils, are often produced. I have seen a leaflet low down in the compound leaf ofVicia sativaconverted into a tendril, and a tendril possesses many peculiar properties, such as spontaneous movement and irritability. The calyx sometimes assumes, either wholly or by stripes, the colour and texture of the corolla. Stamens are so frequently converted, more or less completely, into petals, that such cases are passed over as not deserving notice; but as petals have special functions to perform, namely, to protect the included organs, to attract insects, and in not a few cases to guide their entrance by well-adapted contrivances, we can hardly account for the conversion of stamens into petals merely by unnatural or superfluous nourishment. Again, the edge of a petal may occasionally be found including one of the highest products of the plant, namely the pollen; for instance, I have seen in an Ophrys a pollen-mass with its curious structure of little packets, united together and to the caudicle by elastic threads, formed between the edges of an upper petal. The segments of the calyx of the common pea have been observed partially converted into carpels, including ovules, and with their tips converted into stigmas. Numerous analogous facts could be given.[923]
I do not know how physiologists look at such facts as the foregoing. According to the doctrine of pangenesis, the free and superabundant gemmules of the transposed organs are developed in the wrong place, from uniting with wrong cells or aggregates of cells during their nascent state; and this would follow from a slight modification in the elective affinity of such cells, or possibly of certain gemmules. Nor ought we to feel much surprise at the affinities of cells and gemmules varyingunder domestication, when we remember the many curious cases given, in the seventeenth chapter, of cultivated plants which absolutely refuse to be fertilised by their own pollen or by that of the same species, but are abundantly fertile with pollen of a distinct species; for this implies that their sexual elective affinities—and this is the term used by Gärtner—have been modified. As the cells of adjoining or homologous parts will have nearly the same nature, they will be liable to acquire by variation each other's elective affinities; and we can thus to a certain extent understand such cases as a crowd of horns on the heads in certain sheep, of several spurs on the leg, and of hackles on the head of the fowl, and with the pigeon the occurrence of wing-feathers on their legs and of membrane between their toes; for the leg is the homologue of the wing. As all the organs of plants are homologous and spring from a common axis, it is natural that they should be eminently liable to transposition. It ought to be observed that when any compound part, such as an additional limb or an antenna, springs from a false position, it is only necessary that the few first gemmules should be wrongly attached; for these whilst developing would attract others in due succession, as in the regrowth of an amputated limb. When parts which are homologous and similar in structure, as the vertebræ in snakes or the stamens in polyandrous flowers, &c., are repeated many times in the same organism, closely allied gemmules must be extremely numerous, as well as the points to which they ought to become united; and, in accordance with the foregoing views, we can to a certain extent understand Isid. Geoffroy St. Hilaire's law, namely, that parts, which are already multiple, are extremely liable to vary in number.
The same general principles apply to the fusion of homologous parts; and with respect to mere cohesion there is probably always some degree of fusion, at least near the surface. When two embryos during their early development come into close contact, as both include corresponding gemmules, which must be in all respects almost identical in nature, it is not surprising that some derived from one embryo and some from the other should unite at the point of contact with a single nascent cell or aggregate of cells, and thus give rise to a single part or organ. For instance, two embryos might thus come to have on theiradjoining sides a single symmetrical arm, which in one sense will have been formed by the fusion of the bones, muscles, &c., belonging to the arms of both embryos. In the case of the fish described by Lereboullet, in which a double head was seen gradually to fuse into a single one, the same process must have taken place, together with the absorption of all the parts which had been already formed. These cases are exactly the reverse of those in which a part is doubled either spontaneously or after an injury; for in the case of doubling, the superabundant gemmules of the same part are separately developed in union with adjoining points; whilst in the case of fusion the gemmules derived from two homologous parts become mingled and form a single part; or it may be that the gemmules from one of two adjoining embryos alone become developed.
Variability often depends, as I have attempted to show, on the reproductive organs being injuriously affected by changed conditions; and in this case the gemmules derived from the various parts of the body are probably aggregated in an irregular manner, some superfluous and others deficient. Whether a superabundance of gemmules, together with fusion during development, would lead to the increased size of any part cannot be told; but we can see that their partial deficiency, without necessarily leading to the entire abortion of the part, might cause considerable modifications; for in the same manner as a plant, if its own pollen be excluded, is easily hybridised, so, in the case of a cell, if the properly succeeding gemmules were absent, it would probably combine easily with other and allied gemmules. We see this in the case of imperfect nails growing on the stumps of amputated fingers,[924]for the gemmules of the nails have manifestly been developed at the nearest point.
In variations caused by the direct action of changed conditions, whether of a definite or indefinite nature, as with the fleeces of sheep in hot countries, with maize grown in cold countries, with inherited gout, &c., the tissues of the body, according to the doctrine of pangenesis, are directly affected by the new conditions, and consequently throw off modified gemmules, which are transmitted with their newly acquired peculiarities to the offspring. On any ordinary view it is unintelligible how changedconditions, whether acting on the embryo, the young or adult animal, can cause inherited modifications. It is equally or even more unintelligible on any ordinary view, how the effects of the long-continued use or disuse of any part, or of changed habits of body or mind, can be inherited. A more perplexing problem can hardly be proposed; but on our view we have only to suppose that certain cells become at last not only functionally but structurally modified; and that these throw off similarly modified gemmules. This may occur at any period of development, and the modification will be inherited at a corresponding period; for the modified gemmules will unite in all ordinary cases with the proper preceding cells, and they will consequently be developed at the same period at which the modification first arose. With respect to mental habits or instincts, we are so profoundly ignorant on the relation between the brain and the power of thought that we do not know whether an inveterate habit or trick induces any change in the nervous system; but when any habit or other mental attribute, or insanity, is inherited, we must believe that some actual modification is transmitted;[925]and this implies, according to our hypothesis, that gemmules derived from modified nerve-cells are transmitted to the offspring.
It is generally, perhaps always, necessary that an organism should be exposed during several generations to changed conditions or habits, in order that any modification in the structure of the offspring should ensue. This may be partly due to the changes not being at first marked enough to catch the attention, but this explanation is insufficient; and I can account for the fact, only by the assumption, which we shall see under the head of reversion is strongly supported, that gemmules derived from each cell before it had undergone the least modification are transmitted in large numbers to successive generations, but that the gemmules derived from the same cells after modification, naturally go on increasing under the same favouring conditions, until at last they become sufficiently numerous to overpower and supplant the old gemmules.
Another difficulty may be here noticed; we have seen thatthere is an important difference in the frequency, though not in the nature, of the variations in plants propagated by sexual and asexual generation. As far as variability depends on the imperfect action of the reproductive organs under changed conditions, we can at once see why seedlings should be far more variable than plants propagated by buds. We know that extremely slight causes,—for instance, whether a tree has been grafted or grows on its own stock, the position of the seeds within the capsule, and of the flowers on the spike,—sometimes suffice to determine the variation of a plant, when raised from seed. Now, it is probable, as explained when discussing alternate generation, that a bud is formed of a portion of already differentiated tissue; consequently an organism thus formed does not pass through the earlier phases of development, and cannot be so freely exposed, at the age when its structure would be most readily modified, to the various causes inducing variability; but it is very doubtful whether this is a sufficient explanation of the difficulty.
With respect to the tendency to reversion, there is a similar difference between plants propagated from buds and seed. Many varieties, whether originally produced from seed or buds, can be securely propagated by buds, but generally or invariably revert by seed. So, also, hybridised plants can be multiplied to any extent by buds, but are continually liable to reversion by seed,—that is, to the loss of their hybrid or intermediate character. I can offer no satisfactory explanation of this fact. Here is a still more perplexing case: certain plants with variegated leaves, phloxes with striped flowers, barberries with seedless fruit, can all be securely propagated by the buds on cuttings; but the buds developed from the roots of these cuttings almost invariably lose their character and revert to their former condition.
Finally, we can see on the hypothesis of pangenesis that variability depends on at least two distinct groups of causes. Firstly, on the deficiency, superabundance, fusion, and transposition of gemmules, and on the redevelopment of those which have long been dormant. In these cases the gemmules themselves have undergone no modification; but the mutations in the above respects will amply account for much fluctuatingvariability. Secondly, in the cases in which the organisation has been modified by changed conditions, the increased use or disuse of parts, or any other cause, the gemmules cast off from the modified units of the body will be themselves modified, and, when sufficiently multiplied, will be developed into new and changed structures.
Turning now to Inheritance: if we suppose a homogeneous gelatinous protozoon to vary and assume a reddish colour, a minute separated atom we aid naturally, as it grew to full size, retain the same colour; and we should have the simplest form of inheritance.[926]Precisely the same view may be extended to the infinitely numerous and diversified units of which the whole body in one of the higher animals is composed; and the separated atoms are our gemmules. We have already sufficiently discussed the inheritance of the direct effects of changed conditions, and of increased use or disuse of parts, and, by implication, the important principle of inheritance at corresponding ages. These groups of facts are to a large extent intelligible on the hypothesis of pangenesis, and on no other hypothesis as yet advanced.
A few words must be added on the complete abortion or suppression of organs. When a part becomes diminished by disuse prolonged during many generations, the principle of economy of growth, as previously explained, will tend to reduce it still further; but this will not account for the complete or almost complete obliteration of, for instance, a minute papilla of cellular tissue representing a pistil, or of a microscopically minute nodule of bone representing a tooth. In certain cases of suppression not yet completed, in which a rudiment occasionally reappears through reversion, diffused gemmules derived from this part must, according to our view, still exist; hence we must suppose that the cells, in union with which the rudiment was formerly developed, in these cases fail in their affinity for such gemmules. But in the cases of complete and final abortion the gemmules themselves no doubt have perished; nor is thisin any way improbable, for, though a vast number of active and long-dormant gemmules are diffused and nourished in each living creature, yet there must be some limit to their number; and it appears natural that gemmules derived from an enfeebled and useless rudiment would be more liable to perish than those derived from other parts which are still in full functional activity.
With respect to mutilations, it is certain that a part may be removed or injured during many generations, and no inherited result follow; and this is an apparent objection to the hypothesis which will occur to every one. But, in the first place, a being can hardly be intentionally mutilated during its early stages of growth whilst in the womb or egg; and such mutilations, when naturally caused, would appear like congenital deficiencies, which are occasionally inherited. In the second place, according to our hypothesis, gemmules multiply by self-division and are transmitted from generation to generation; so that during a long period they would be present and ready to reproduce a part which was repeatedly amputated. Nevertheless it appears, from the facts given in the twelfth chapter, that in some rare cases mutilations have been inherited, but in most of these the mutilated surface became diseased. In this case it may be conjectured that the gemmules of the lost part were gradually all attracted by the partially diseased surface, and thus perished. Although this would occur in the injured individual alone, and therefore in only one parent, yet this might suffice for the inheritance of a mutilation, on the same principle that a hornless animal of either sex, when crossed with a perfect animal of the opposite sex, often transmits its deficiency.
The last subject that need here be discussed, namely Reversion, rests on the principle that transmission and development, though generally acting in conjunction, are distinct powers; and the transmission of gemmules and their subsequent development show us how the existence of these two distinct powers is possible. We plainly see this distinction in the many cases in which a grandfather transmits to his grandson, through his daughter, characters which she does not, or cannot, possess. Why the development of certain characters, not necessarily in any way connected with the reproductive organs, should be confined to one sex alone—that is, why certain cells in one sexshould unite with and cause the development of certain gemmules—we do not in the least know; but it is the common attribute of most organic beings in which the sexes are separate.
The distinction between transmission and development is likewise seen in all ordinary cases of Reversion; but before discussing this subject it may be advisable to say a few words on those characters which I have called latent, and which would not be classed under Reversion in its usual sense. Most, or perhaps all, the secondary characters, which appertain to one sex, lie dormant in the other sex; that is, gemmules capable of development into the secondary male sexual characters are included within the female; and conversely female characters in the male. Why in the female, when her ovaria become diseased or fail to act, certain masculine gemmules become developed, we do not clearly know, any more than why when a young bull is castrated his horns continue growing until they almost resemble those of a cow; or why, when a stag is castrated, the gemmules derived from the antlers of his progenitors quite fail to be developed. But in many cases, with variable organic beings, the mutual affinities of the cells and gemmules become modified, so that parts are transposed or multiplied; and it would appear that a slight change in the constitution of an animal, in connection with the state of the reproductive organs, leads to changed affinities in the tissues of various parts of the body. Thus, when male animals first arrive at puberty, and subsequently during each recurrent season, certain cells or parts acquire an affinity for certain gemmules, which become developed into the secondary masculine characters; but if the reproductive organs be destroyed, or even temporarily disturbed by changed conditions, these affinities are not excited. Nevertheless, the male, before he arrives at puberty, and during the season when the species does not breed, must include the proper gemmules in a latent state. The curious case formerly given of a Hen which assumed the masculine characters, not of her own breed but of a remote progenitor, illustrates the connexion between latent sexual characters and ordinary reversion. With those animals and plants which habitually produce several forms, as with certain butterflies described by Mr. Wallace, in which three female forms andthe male exist, or as with the trimorphic species of Lythrum and Oxalis, gemmules capable of reproducing several widely-different forms must be latent in each individual.
The same principle of the latency of certain characters, combined with the transposition of organs, may be applied to those singular cases of butterflies and other insects, in which exactly one half or one quarter of the body resembles the male, and the other half or three quarters the female; and when this occurs the opposite sides of the body, separated from each other by a distinct line, sometimes differ in the most conspicuous manner. Again, these same principles apply to the cases given in the thirteenth chapter, in which the right and left sides of the body differ to an extraordinary degree, as in the spiral winding of certain shells, and as in the genus Verruca among cirripedes; for in these cases it is known that either side indifferently may undergo the same remarkable change of development.
Reversion, in the ordinary sense of the word, comes into action so incessantly, that it evidently forms an essential part of the general law of inheritance. It occurs with beings, however propagated, whether by buds or seminal generation, and sometimes may even be observed in the same individual as it advances in age. The tendency to reversion is often induced by a change of conditions, and in the plainest manner by the act of crossing. Crossed forms are generally at first nearly intermediate in character between their two parents; but in the next generation the offspring generally revert to one or both of their grandparents, and occasionally to more remote ancestors. How can we account for these facts? Each organic unit in a hybrid must throw off, according to the doctrine of pangenesis, an abundance of hybridised gemmules, for crossed plants can be readily and largely propagated by buds; but by the same hypothesis there will likewise be present dormant gemmules derived from both pure parent-forms; and as these latter retain their normal condition, they would, it is probable, be enabled to multiply largely during the lifetime of each hybrid. Consequently the sexual elements of a hybrid will include both pure and hybridised gemmules; and when two hybrids pair, the combination of pure gemmules derived from the one hybrid with the pure gemmules of the same parts derived from the other wouldnecessarily lead to complete reversion of character; and it is, perhaps, not too bold a supposition that unmodified and undeteriorated gemmules of the same nature would be especially apt to combine. Pure gemmules in combination with hybridised gemmules would lead to partial reversion. And lastly, hybridised gemmules derived from both parent-hybrids would simply reproduce the original hybrid form.[927]All these cases and degrees of reversion incessantly occur.