The Variation of Animals and Plants under Domestication — Volume 2

It is well known from innumerable trials made in all parts of the world, that buds may be inserted into a stock, and that the plants thus raised are not affected in a greater degree than can be accounted for by changed nutrition. Nor do the seedlings raised from such inserted buds partake of the character of the stock, though they are more liable to vary than are seedlings from the same variety growing on its own roots. A bud, also, may sport into a new and strongly-marked variety without any other bud on the same plant being in the least degree affected. We may therefore infer, in accordance with the common view, that each bud is a distinct individual, and that its formative elements do not spread beyond the parts subsequently developed from it. Nevertheless, we have seen in the abstract on graft-hybridisation in the eleventh chapter that buds certainly include formative matter, which can occasionally combine with that included in the tissues of a distinct variety or species; a plant intermediate between the two parent-forms being thus produced. In the case of the potato we have seen that the tubers produced from a bud of one kind inserted into another are intermediate in colour, size, shape and state of surface; that the stems, foliage, and even certain constitutional peculiarities, such as precocity, are likewise intermediate. With these well- established cases, the evidence that graft-hybrids have also been produced with the laburnum, orange, vine, rose, etc., seems sufficient. But we do not know under what conditions this rare form of reproduction is possible. From these several cases we learn the important fact that formative elements capable of blending with those of a distinct individual (and this is the chief characteristic of sexual generation), are not confined to the reproductive organs, but are present in the buds and cellular tissue of plants; and this is a fact of the highest physiological importance.

DIRECT ACTION OF THE MALE ELEMENT ON THE FEMALE.

In the eleventh chapter, abundant proofs were given that foreign pollen occasionally affects in a direct manner the mother-plant. Thus, when Gallesio fertilised an orange-flower with pollen from the lemon, the fruit bore stripes of perfectly characterised lemon-peel. With peas, several observers have seen the colour of the seed-coats and even of the pod directly affected by the pollen of a distinct variety. So it has been with the fruit of the apple, which consists of the modified calyx and upper part of the flower-stalk. In ordinary cases these parts are wholly formed by the mother-plant. We here see that the formative elements included within the male element or pollen of one variety can affect and hybridise, not the part which they are properly adapted to affect, namely, the ovules, but the partially-developed tissues of a distinct variety or species. We are thus brought half-way towards a graft- hybrid, in which the formative elements included within the tissues of one individual combine with those included in the tissues of a distinct variety or species, thus giving rise to a new and intermediate form, independently of the male or female sexual organs.

With animals which do not breed until nearly mature, and of which all the parts are then fully developed, it is hardly possible that the male element should directly affect the female. But we have the analogous and perfectly well-ascertained case of the male element affecting (as with the quagga and Lord Morton's mare) the female or her ova, in such a manner that when she is impregnated by another male her offspring are affected and hybridised by the first male. The explanation would be simple if the spermatozoa could keep alive within the body of the female during the long interval which has sometimes elapsed between the two acts of impregnation; but no one will suppose that this is possible with the higher animals.

DEVELOPMENT.

The fertilised germ reaches maturity by a vast number of changes: these are either slight and slowly effected, as when the child grows into the man, or are great and sudden, as with the metamorphoses of most insects. Between these extremes we have every gradation, even within the same class; thus, as Sir J. Lubbock has shown (27/28. 'Transact. Linn. Soc.' volume 24 1863 page 62.) there is an Ephemerous insect which moults above twenty times, undergoing each time a slight but decided change of structure; and these changes, as he further remarks, probably reveal to us the normal stages of development, which are concealed and hurried through or suppressed in most other insects. In ordinary metamorphoses, the parts and organs appear to become changed into the corresponding parts in the next stage of development; but there is another form of development, which has been called by Professor Owen metagenesis. In this case "the new parts are not moulded upon the inner surface of the old ones. The plastic force has changed its course of operation. The outer case, and all that gave form and character to the precedent individual, perish and are cast off; they are not changed into the corresponding parts of the new individual. These are due to a new and distinct developmental process," etc. (27/29. 'Parthenogenesis' 1849 pages 25, 26. Prof. Huxley has some excellent remarks ('Medical Times' 1856 page 637) on this subject in reference to the development of star-fishes, and shows how curiously metamorphosis graduates into gemmation or zoid-formation, which is in fact the same as metagenesis.) Metamorphosis, however, graduates so insensibly, into metagenesis, that the two processes cannot be distinctly separated. For instance, in the last change which Cirripedes undergo, the alimentary canal and some other organs are moulded on pre-existing parts; but the eyes of the old and the young animal are developed in entirely different parts of the body; the tips of the mature limbs are formed within the larval limbs, and may be said to be metamorphosed from them; but their basal portions and the whole thorax are developed in a plane at right angles to the larval limbs and thorax; and this may be called metagenesis. The metagenetic process is carried to an extreme point in the development of some Echinoderms, for the animal in the second stage of development is formed almost like a bud within the animal of the first stage, the latter being then cast off like an old vestment, yet sometimes maintaining for a short period an independent vitality. (27/30. Prof. J. Reay Greene in Gunther's 'Record of Zoolog. Lit.' 1865 page 625.) If, instead of a single individual, several were to be thus developed metagenetically within a pre- existing form, the process would be called one of alternate generation. The young thus developed may either closely resemble the encasing parent-form, as with the larvae of Cecidomyia, or may differ to an astonishing degree, as with many parasitic worms and jelly-fishes; but this does not make any essential difference in the process, any more than the greatness or abruptness of the change in the metamorphoses of insects.

The whole question of development is of great importance for our present subject. When an organ, the eye, for instance, is metagenetically formed in a part of the body where during the previous stage of development no eye existed, we must look at it as a new and independent growth. The absolute independence of new and old structures, although corresponding in structure and function, is still more obvious when several individuals are formed within a previous form, as in the cases of alternate generation. The same important principle probably comes largely into play even in the case of apparently continuous growth, as we shall see when we consider the inheritance of modifications at corresponding ages.

We are led to the same conclusion, namely, the independence of parts successively developed, by another and quite distinct group of facts. It is well known that many animals belonging to the same order, and therefore not differing widely from each other, pass through an extremely different course of development. Thus certain beetles, not in any way remarkably different from others of the same order, undergo what has been called a hyper-metamorphosis — that is, they pass through an early stage wholly different from the ordinary grub-like larva. In the same sub-order of crabs, namely, the Macroura, as Fritz Muller remarks, the river cray-fish is hatched under the same form which it ever afterwards retains; the young lobster has divided legs, like a Mysis; the Palaemon appears under the form of a Zoea, and Peneus under the Nauplius- form; and how wonderfully these larval forms differ from one another, is known to every naturalist. (27/31. Fritz Muller 'Fur Darwin' 1864 s. 65, 71. The highest authority on crustaceans, Prof. Milne-Edwards, insists ('Annal. des Sci. Nat.' 2nd series Zoolog. tome 3 page 322) on the difference in the metamorphosis of closely-allied genera.) Some other crustaceans, as the same author observes, start from the same point and arrive at nearly the same end, but in the middle of their development are widely different from one another. Still more striking cases could be given with respect to the Echinodermata. With the Medusae or jelly-fishes Professor Allman observes, "The classification of the Hydroida would be a comparatively simple task if, as has been erroneously asserted, generically-identical medusoids always arose from generically-identical polypoids; and, on the other hand, that generically- identical polypoids always gave origin to generically-identical medusoids." So again, Dr. Strethill Wright remarks, "In the life-history of the Hydroidae any phase, planuloid, polypoid, or medusoid, may be absent." (27/32. Prof. Allman 'Annals and Mag. of Nat. Hist.' 3rd series volume 13 1864 page 348; Dr. S. Wright ibid volume 8 1861 page 127. See also page 358 for analogous statements by Sars.)

According to the belief now generally accepted by our best naturalists, all the members of the same order or class, for instance, the Medusae or the Macrourous crustaceans, are descended from a common progenitor. During their descent they have diverged much in structure, but have retained much in common; and this has occurred, though they have passed through and still pass through marvellously different metamorphoses. This fact well illustrates how independent each structure is from that which precedes and that which 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 one another. Each organ, says Claude Bernard (27/33. 'Tissus Vivants' 1866 page 22.), has its proper life, its autonomy; it can develop and reproduce itself independently of the adjoining tissues. A great German authority, Virchow (27/34. 'Cellular Pathology' translated by Dr. Chance 1860 pages 14, 18, 83, 460.), asserts still more emphatically that each system 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 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 Sir J. Paget remarks, lives its appointed time and then dies, and is replaced after being cast off or absorbed. (27/35. Paget 'Surgical Pathology' volume 1 1853 pages 12-14.) 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 Sir J. Paget (27/36. Ibid page 19.) 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 ear of an ox, lived for eight years, and acquired a weight of 396 grammes (nearly fourteen ounces), and the astonishing length of twenty-four centimetres, or about nine inches; so that the head of the ox appeared to bear three horns. (27/37. See Prof. Mantegazza's interesting work 'Degli innesti Animali' etc. Milano 1865 page 51 tab. 3.) The tail of a pig has been grafted into the middle of its back, and reacquired sensibility. Dr. Ollier (27/38. 'De la Production Artificielle des Os' page 8.) 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. The frequent presence of hairs and of perfectly developed teeth, even teeth of the second dentition, in ovarian tumours (27/39. Isidore Geoffroy Saint-Hilaire 'Hist. des Anomalies' tome 2 pages 549, 560, 562; Virchow ibid page 484. Lawson Tait 'The Pathology of Diseases of the Ovaries' 1874 pages 61, 62.), are facts leading to the same conclusion. Mr. Lawson Tait refers to a tumour in which "over 300 teeth were found, resembling in many respects milk-teeth;" and to another tumour, "full of hair which had grown and been shed from one little spot of skin not bigger than the tip of my little finger. The amount of hair in the sac, had it grown from a similarly sized area of the scalp, would have taken almost a lifetime to grow and be shed."

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. (27/40. For the most recent classification of cells, see Ernst Hackel 'Generelle Morpholog.' b. 2 1866 s. 275.) The doctrine of omnis cellula e cellula is admitted for plants, and widely prevails with respect to animals. (27/41. Dr. W. Turner 'The Present Aspect of Cellular Pathology' 'Edinburgh Medical Journal' April 1863.) 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 every one. But when an organism undergoes great changes 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 attributed by the supporters of the cellular doctrine to some inherent power which the cells possess, and not to any external agency. Others maintain that cells and tissues of all kinds may be formed, independently of pre-existing cells, from plastic lymph or blastema. Whichever view may be correct, every one admits that the body consists of a multitude of organic units, all of which possess their own proper attributes, and are 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. The fluctuating variability thus induced is apparently due in part to the sexual system being easily affected, 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 in the cases of bud-variation. Although we are seldom able to trace the nature of the connection, many deviations of structure no doubt result from changed conditions acting directly on the organisation, independently of the reproductive system. 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, of which several instances have been given. But it is by no means clear why the offspring should be affected by the exposure of the parents to new conditions, and why it is necessary in most cases that several generations should have been thus exposed.

How, again, can we explain 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 diminished and increased in a corresponding manner 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 these mental 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.

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; and we may on the whole conclude that 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.

Inheritance is governed by various laws. Characters which first appear at any particular age tend to reappear at a corresponding age. They often become associated with certain seasons of the year, and reappear in the offspring at a corresponding season. If they appear rather late in life in one sex, they tend to reappear exclusively in the same sex at the same period of life.

The principle of reversion, recently alluded to, is one of the most wonderful of 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 that changed conditions have the power of evoking long-lost characters, as in the case of animals becoming feral. 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, both 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 of the species, but of which not even a rudiment is left, suddenly reappears, as with the fifth stamen in some Scrophulariaceae. 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, if of crossed parentage, — as in the rare cases described of fowls, pigeons, cattle, and rabbits, which have reverted to the colours of one of their parents or ancestors as they advanced in years.

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 the secondary characters of the opposite sex lie latent and ready to be evolved when the reproductive organs are injured. This comparison of the secondary sexual characters which lie latent in both sexes, with other latent characters, is the more appropriate from the case recorded of a 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. In every living creature we may feel assured that a host of long-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 see connected by some intelligible bond. This can be done, if we make the following assumptions, and much may be advanced in favour of the chief one. The secondary assumptions can likewise be supported by various physiological considerations. It is universally admitted that the cells or units of the body increase by self-division or proliferation, retaining the same nature, and that they ultimately become converted into the various tissues and substances of the body. But besides this means of increase I assume that the units throw off minute granules which are dispersed throughout the whole system; that these, when supplied with proper nutriment, multiply by self-division, and are ultimately developed into units like those from which they were originally derived. These granules may be called gemmules. They are collected from all parts of the system to constitute the sexual elements, and their development in the next generation forms a new being; but they are likewise capable of transmission in a dormant state to future generations and may then be developed. Their development depends on their union with other partially developed or nascent cells 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 unit, not only during the adult state, but during each stage of development of every organism; but not necessarily during the continued existence of the same unit. Lastly, I assume that the gemmules in their dormant state have a mutual affinity for each other, leading to their aggregation into buds or into the sexual elements. Hence, it is not the reproductive organs or buds which generate new organisms, but the units of which each individual is composed. These assumptions constitute the provisional hypothesis which I have called Pangenesis. Views in many respects similar have been propounded by various authors. (27/42. Mr. G.H. Lewes ('Fortnightly Review' November 1, 1868 page 506) remarks on the number of writers who have advanced nearly similar views. More than two thousand years ago Aristotle combated a view of this kind, which, as I hear from Dr. W. Ogle, was held by Hippocrates and others. Ray, in his 'Wisdom of God' (2nd edition 1692 page 68), says that "every part of the body seems to club and contribute to the seed." The "organic molecules" of Buffon ('Hist. Nat. Gen.' edition of 1749 tome 2 pages 54, 62, 329, 333, 420, 425) appear at first sight to be the same as the gemmules of my hypothesis, but they are essentially different. Bonnet ('Oeuvres d'Hist. Nat.' tome 5 part 1 1781 4to edition page 334) speaks of the limbs having germs adapted for the reparation of all possible losses; but whether these germs are supposed to be the same with those within buds and the sexual organs is not clear. Prof. Owen says ('Anatomy of Vertebrates' volume 3 1868 page 813) that he fails to see any fundamental difference between the views which he propounded in his 'Parthenogenesis' (1849 pages 5- 8), and which he now considers as erroneous, and my hypothesis of pangenesis: but a reviewer ('Journal of Anat. and Phys.' May 1869 page 441) shows how different they really are. I formerly thought that the "physiological units" of Herbert Spencer ('Principles of Biology' volume 1 chapters 4 and 8 1863-64) were the same as my gemmules, but I now know that this is not the case. Lastly, it appears from a review of the present work by Prof. Mantegazza ('Nuova Antologia, Maggio' 1868), that he (in his 'Elementi di Igiene' Ediz. 3 page 540) clearly foresaw the doctrine of pangenesis.)

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, as simple as possible, of the hypothesis. If one of the Protozoa be formed, as it appears under the microscope, of a small mass of homogeneous gelatinous matter, a minute particle or gemmule thrown off from any part and nourished under favourable circumstances would reproduce the whole; but if the upper and lower surfaces were to differ in texture from each other and from the central portion, then all three parts would have to throw off gemmules, which when aggregated by mutual affinity would form either buds or the sexual elements, and would ultimately be developed into a similar organism. 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 at each stage of development; these gemmules being developed in union with pre-existing nascent cells in due order of succession.

Physiologists maintain, as we have seen, that each unit of the body, though to a large extent dependent on others, is likewise to a certain extent independent or autonomous, and has the power of increasing by self-division. I go one step further, and assume that each unit casts off free gemmules which are dispersed throughout the system, and are capable under proper conditions of being developed into similar units. Nor can this assumption be considered as gratuitous and improbable. It is manifest that the sexual elements and buds include formative matter of some kind, capable of development; and we now know from the production of graft-hybrids that similar matter is dispersed throughout the tissues of plants, and can combine with that of another and distinct plant, giving rise to a new being, intermediate in character. We know also that the male element can act directly on the partially developed tissues of the mother-plant, and on the future progeny of female animals. The formative matter which is thus dispersed throughout the tissues of plants, and which is capable of being developed into each unit or part, must be generated there by some means; and my chief assumption is that this matter consists of minute particles or gemmules cast off from each unit or cell. (27/43. Mr. Lowne has observed ('Journal of Queckett Microscopical Club' September 23, 1870) certain remarkable changes in the tissues of the larva of a fly, which makes him believe "it possible that organs and organisms are sometimes developed by the aggregation of excessively minute gemmules, such as those which Mr. Darwin's hypothesis demands.")

But I have further to assume that the gemmules in their undeveloped state are capable of largely multiplying themselves by self-division, like independent organisms. Delpino insists that to "admit of multiplication by fissiparity in corpuscles, analogous to seeds or buds...is repugnant to all analogy." But this seems a strange objection, as Thuret (27/44. 'Annales des Sc. Nat.' 3rd series Bot. tome 14 1850 page 244.) has seen the zoospore of an alga divide itself, and each half germinated. Haeckel divided the segmented ovum of a siphonophora into many pieces, and these were developed. Nor does the extreme minuteness of the gemmules, which can hardly differ much in nature from the lowest and simplest organisms, render it improbable that they should grow and multiply. A great authority, Dr. Beale (27/45. 'Disease Germs' page 20.), says "that minute yeast cells are capable of throwing off buds or gemmules, much less than the 1/100000 of an inch in diameter;" and these he thinks are "capable of subdivision practically ad infinitum."

A particle of small-pox matter, so minute as to be borne by the wind, must multiply itself many thousandfold in a person thus inoculated; and so with the contagious matter of scarlet fever. (27/46. See some very interesting papers on this subject by Dr. Beale in 'Medical Times and Gazette' September 9, 1865 pages 273, 330.) It has recently been ascertained (27/47. Third Report of the R. Comm. on the Cattle Plague as quoted in 'Gardener's Chronicle' 1866 page 446.) 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 will 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 will appear still more improbable; but here again we should remember that many rudimentary and useless organs 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 ovule, and within each spermatozoon or pollen-grain, and as some animals and plants produce an astonishing number of pollen-grains and ovules (27/48. Mr. F. Buckland found 6,867,840 eggs in a cod-fish ('Land and Water' 1868 page 62). An Ascaris produces about 64,000,000 eggs (Carpenter's 'Comp. Phys.' 1854 page 590). Mr. J. Scott, of the Royal Botanic Garden of Edinburgh, calculated, in the same manner as I have done for some British Orchids ('Fertilisation of Orchids' page 344), the number of seeds in a capsule of an Acropera and found the number to be 371,250. Now this plant produces several flowers on a raceme, and many racemes during a season. In an allied genus, Gongora, Mr. Scott has seen twenty capsules produced on a single raceme; ten such racemes on the Acropera would yield above seventy-four millions of seed.), the number and minuteness of the gemmules must be something inconceivable. But considering how minute the molecules are, and how many go to the formation of the smallest granule of any ordinary substance, this difficulty with respect to the gemmules is not insuperable. From the data arrived at by Sir W. Thomson, my son George finds that a cube of 1/10000 of an inch of glass or water must consist of between 16 million millions, and 131 thousand million million molecules. No doubt the molecules of which an organism is formed are larger, from being more complex, than those of an inorganic substance, and probably many molecules go to the formation of a gemmule; but when we bear in mind that a cube of 1/10000 of an inch is much smaller than any pollen-grain, ovule or bud, we can see what a vast number of gemmules one of these bodies might contain.

The gemmules derived from each part or organ must be thoroughly dispersed throughout the whole system. We know, for instance, that even a minute fragment of a leaf of a Begonia will reproduce the whole plant; and that if a fresh-water worm is chopped into small pieces, each will reproduce the whole animal. Considering also the minuteness of the gemmules and the permeability of all organic tissues, the thorough dispersion of the gemmules is not surprising. That matter may be readily transferred without the aid of vessels from part to part of the body, we have a good instance in a case recorded by Sir J. Paget of a lady, whose hair lost its colour at each successive attack of neuralgia and recovered it again in the course of a few days. With plants, however, and probably with compound animals, such as corals, the gemmules do not ordinarily spread from bud to bud, but are confined to the parts developed from each separate bud; and of this fact no explanation can be given.

The assumed elective affinity of each gemmule for that particular cell which precedes it in due order of development is supported by many analogies. In all ordinary cases of sexual reproduction, the male and female elements certainly have a mutual affinity for each other: thus, it is believed that about ten thousand species of Compositae 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 formative matter of each part contained within the ovules and the male element act on each other by some law of special affinity, so that corresponding parts affect one another; thus, a calf produced from a short-horned cow by a long-horned bull has its horns affected by the union of the two forms, and the offspring from two birds with differently coloured tails have their tails affected.

The various tissues of the body plainly show, as many physiologists have insisted (27/49. Paget 'Lectures on Pathology' page 27; Virchow 'Cellular Pathology' translated by Dr. Chance pages 123, 126, 294. Claude Bernard 'Des Tissus Vivants' pages 177, 210, 337; Muller 'Physiology' English translation page 290.), 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 curare affecting certain nerves; Lytta vesicatoria the kidneys; and the poisonous matter of various diseases, as small-pox, scarlet- fever, hooping-cough, glanders, and hydrophobia, affecting certain definite parts of the body. 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 precedes it in due order of growth. That the formative matter within the pollen of plants, which by our hypothesis consists of gemmules, can unite with and modify the partially developed cells of the mother-plant, we have clearly seen in the section devoted to this subject. As the tissues of plants are formed, as far as is known, only by the proliferation of pre-existing cells, we must conclude that the gemmules derived from the foreign pollen do not become developed into new and separate cells, but penetrate and modify the nascent cells of the mother-plant. This process may be compared with what takes place in the act of ordinary fertilisation, during which the contents of the pollen-tubes penetrate the closed embryonic sac 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. In this case and in all others the proper gemmules must combine in due order with pre-existing nascent cells, owing to their elective affinities. A slight difference in nature between the gemmules and the nascent cells would be far from interfering with their mutual union and development, for we well know in the case of ordinary reproduction that such slight differentiation in the sexual elements favours in a marked manner their union and subsequent development, as well as the vigour of the offspring thus produced.

Thus far we have been able by the aid of our hypothesis to throw some obscure light on the problems which have come before us; but it must be confessed that many points remain altogether doubtful. Thus it is useless to speculate at what period of development each unit of the body casts off its gemmules, as the whole subject of the development of the various tissues is as yet far from clear. We do not know whether the gemmules are merely collected by some unknown means at certain seasons within the reproductive organs, or whether after being thus collected they rapidly multiply there, as the flow of blood to these organs at each breeding season seems to render probable. Nor do we know why the gemmules collect to form buds in certain definite places, leading to the symmetrical growth of trees and corals. We have no means of deciding whether the ordinary wear and tear of the tissues is made good by means of gemmules, or merely by the proliferation of pre-existing cells. If the gemmules are thus consumed, as seems probable from the intimate connection between the repair of waste, regrowth, and development, and more especially from the periodical changes which many male animals undergo in colour and structure, then some light would be thrown on the phenomena of old age, with its lessened power of reproduction and of the repair of injuries, and on the obscure subject of longevity. The fact of castrated animals, which do not cast off innumerable gemmules in the act of reproduction, not being longer-lived than perfect males, seems opposed to the belief that gemmules are consumed in the ordinary repair of wasted tissues; unless indeed the gemmules after being collected in small numbers within the reproductive organs are there largely multiplied. (27/50. Prof. Ray Lankester has discussed several of the points here referred to as bearing on pangenesis, in his interesting essay, 'On Comparative Longevity in Man and the Lower Animals' 1870 pages 33, 77, etc.)

That the same cells or units may live for a long period and continue multiplying without being modified by their union with free gemmules of any kind, is probable from such cases as that of the spur of a cock which grew to an enormous size when grafted into the ear of an ox. How far units are modified during their normal growth by absorbing peculiar nutriment from the surrounding tissues, independently of their union with gemmules of a distinct nature, is another doubtful point. (27/51. Dr. Ross refers to this subject in his 'Graft Theory of Disease' 1872 page 53.) We shall appreciate this difficulty by calling to mind what complex yet symmetrical growths the cells of plants yield when inoculated by the poison of a gall-insect. With animals various polypoid excrescences and tumours are generally admitted (27/52. Virchow 'Cellular Pathology' translated by Dr. Chance 1860 pages 60, 162, 245, 441, 454.) 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 (27/53. Ibid pages 412-426.), a whole series of permutations and substitutions. "The cartilage cells may be converted 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.

We have good reason to believe that several gemmules are requisite for the development of one and the same unit or cell; for we cannot otherwise understand the insufficiency of a single or even of two or three pollen-grains or spermatozoa. But we are far from knowing whether the gemmules of all the units are free and separate from one another, 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 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 flowers, which are sometimes highly complex structures, with each ridge and hollow contrived for a special purpose, 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 unit. But, as we sometimes see half an anther or a small portion of a filament becoming petali-form, 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 free and separate. Even in so simple a case as that of a perfect cell, with its protoplasmic contents, nucleus, nucleolus, and walls, we do not know whether or not its development depends on a compound gemmule derived from each part. (27/54. See some good criticisms on this head by Delpino and by Mr. G.H. Lewes in the 'Fortnightly Review' November 1, 1868 page 509.)

Having now endeavoured to show that the several foregoing assumptions are to a certain extent supported by analogous facts, and having alluded to 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 one another 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 reversions of the same kind; and as, according to our hypothesis, all the forms of reproduction depend on the aggregation of gemmules derived from the whole body, we can understand this remarkable agreement. Parthenogenesis is no longer wonderful, and if we did not know that great good followed from the union of the sexual elements derived from two distinct individuals, the wonder would be that parthenogenesis did not occur much oftener than it does. On any ordinary theory of reproduction the formation of graft-hybrids, and the action of the male element on the tissues of the mother-plant, as well as on the future progeny of female animals, are great anomalies; but they are intelligible on our hypothesis. The reproductive organs do not actually create the sexual elements; they merely determine the aggregation and perhaps the multiplication of the gemmules in a special manner. These organs, however, together with their accessory parts, have high functions to perform. They adapt one or both elements for independent temporary existence, and for mutual union. The stigmatic secretion acts on the pollen of a plant of the same species in a wholly different manner to what it does on the pollen of one belonging to a distinct genus or family. The spermatophores of the Cephalopoda are wonderfully complex structures, which were formerly mistaken for parasitic worms; and the spermatozoa of some animals possess attributes which, if observed in an independent animal, would be put down to instinct guided by sense-organs, — as when the spermatozoa of an insect find their way into the minute micropyle of the egg.

The antagonism which has long been observed (27/55. Mr. Herbert Spencer ('Principles of Biology' volume 2 page 430) has fully discussed this antagonism.), with certain exceptions, between growth and the power of sexual reproduction (27/56. The male salmon is known to breed at a very early age. The Triton and Siredon, whilst retaining their larval branchiae, according to Filippi and Dumeril ('Annals and Mag. of Nat. Hist.' 3rd series 1866 page 157) are capable of reproduction. Ernst Haeckel has recently ('Monatsbericht Akad. Wiss. Berlin' February 2, 1865) observed the surprising case of a medusa, with its reproductive organs active, which produces by budding a widely different form of medusa; and this latter also has the power of sexual reproduction. Krohn has shown ('Annals and Mag. of Nat. Hist.' 3rd series volume 19 1862 page 6) that certain other medusae, whilst sexually mature, propagate by gemmae. See also Kolliker 'Morphologie und Entwickelungsgeschichte des Pennatulidenstammes' 1872 page 12.) — between the repair of injuries and gemmation — and with plants, between rapid increase by buds, rhizomes, etc., and the production of seed, is partly explained by the gemmules not existing in sufficient numbers for these processes to be carried on simultaneously.

Hardly any fact in physiology is more wonderful than the power of regrowth; for instance, that a snail should be able to reproduce its head, or a salamander its eyes, tail, and legs, exactly at the points where they have been cut off. Such cases are explained by the presence of gemmules derived from each part, and disseminated throughout the body. I have heard the process compared with that of the repair of the broken angles of a crystal by re- crystallisation; 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. But we have here to encounter two objections which apply not only to the regrowth of a part, or of a bisected individual, but to fissiparous generation and budding. The first objection is that the part which is reproduced is in the same stage of development as that of the being which has been operated on or bisected; and in the case of buds, that the new beings thus produced are in the same stage as that of the budding parent. Thus a mature salamander, of which the tail has been cut off, does not reproduce a larval tail; and a crab does not reproduce a larval leg. In the case of budding it was shown in the first part of this chapter that the new being thus produced does not retrograde in development, — that is, does not pass through those earlier stages, which the fertilised germ has to pass through. Nevertheless, the organisms operated on or multiplying themselves by buds must, by our hypothesis, include innumerable gemmules derived from every part or unit of the earlier stages of development; and why do not such gemmules reproduce the amputated part or the whole body at a corresponding early stage of development?

The second objection, which has been insisted on by Delpino, is that the tissues, for instance, of a mature salamander or crab, of which a limb has been removed, are already differentiated and have passed through their whole course of development; and how can such tissues in accordance with our hypothesis attract and combine with the gemmules of the part which is to be reproduced? In answer to these two objections we must bear in mind the evidence which has been advanced, showing that at least in a large number of cases the power of regrowth is a localised faculty, acquired for the sake of repairing special injuries to which each particular creature is liable; and in the case of buds or fissiparous generation, for the sake of quickly multiplying the organism at a period of life when it can be supported in large numbers. These considerations lead us to believe that in all such cases a stock of nascent cells or of partially developed gemmules are retained for this special purpose either locally or throughout the body, ready to combine with the gemmules derived from the cells which come next in due succession. If this be admitted we have a sufficient answer to the above two objections. Anyhow, pangenesis seems to throw a considerable amount of light on the wonderful power of regrowth.

It follows, also, from the view just given, that the sexual elements differ from buds in not including nascent cells or gemmules in a somewhat advanced stage of development, so that only the gemmules belonging to the earliest stages are first developed. As young animals and those which stand low in the scale generally have a much greater capacity for regrowth than older and higher animals, it would also appear that they retain cells in a nascent state, or partially developed gemmules, more readily than do animals which have already passed through a long series of developmental changes. I may here add that although ovules can be detected in most or all female animals at an extremely early age, there is no reason to doubt that gemmules derived from parts modified during maturity can pass into the ovules.

With respect to hybridism, pangenesis agrees well with most of the ascertained facts. We must believe, as previously shown, that several gemmules are requisite for the development of each cell or unit. But from the occurrence of parthenogenesis, more especially from those cases in which an embryo is only partially formed, we may infer that the female element generally includes gemmules in nearly sufficient number for independent development, so that when united with the male element the gemmules are superabundant. Now, when two species or races are crossed reciprocally, the offspring do not commonly differ, and this shows that the sexual elements agree in power, in accordance with the view that both include the same gemmules. Hybrids and mongrels are also 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 on the admission that the gemmules in the fertilised germ are superabundant in number, and that those derived from one parent may 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 occurs in the first generation, or through reversion in succeeding bud and seminal generations, of which fact several instances were given in the eleventh chapter. In these cases we must follow Naudin (27/57. See his excellent discussion on this subject in 'Nouvelles Archives du Museum' tome 1 page 151.) and admit that the "essence" or "element" of the two species, — terms which 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 its characters over the other; and this we can explain by again assuming that the one form has some advantage over the other in the number, vigour, or affinity of its gemmules. In some cases, however, 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. The explanation of this form of prepotency will be obvious when we come to the consideration of Reversion.

When two distinct species are crossed, 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 nascent cells, we need not feel at all surprised that the commixture of gemmules derived from two distinct species should lead to 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 than why unnatural conditions of life, though compatible with health, should cause sterility; or why continued close interbreeding, or the illegitimate unions of heterostyled 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 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.

Each organism reaches maturity through a longer or shorter course of growth and development: the former term being confined to mere increase of size, and development to changed structure. 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 newly formed part may be moulded within a previously existing and corresponding part, and in this case it will appear, falsely as I believe, to be developed from the old part; or it may be formed within a 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 passing through widely different forms; or conversely, after passing through nearly the same early forms, arrive at widely different mature forms. In these cases it is very difficult to accept the common view that the first-formed cells or units possess the inherent power, independently of any external agency, of producing new structures wholly different in form, position, and function. But all these cases become plain on the hypothesis of pangenesis. The 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 becomes partially developed, it unites with (or, to speak metaphorically, is fertilised by) the gemmule of the next succeeding cell, and so onwards. But organisms have often been subjected to changed conditions of life at a certain stage of their development, and in consequence have been slightly modified; and the gemmules cast off from such modified parts will tend to reproduce parts modified in the same manner. This process may be repeated until the structure of the part becomes greatly changed at one particular stage of development, but this will not necessarily affect other parts, whether previously or subsequently formed. In this manner we can understand the remarkable independence of structure in the successive metamorphoses, and especially in the successive metageneses of many animals. 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, we must suppose that the organs were affected at an early age and threw off at this period affected gemmules; but that the affection became visible or injurious only after the prolonged growth, in the strict sense of the word, of the part. In all the changes of structure which regularly supervene during old age, we probably see the effects of deteriorated growth, and not of true development.

The principle of the independent formation of each part, owing to the union of the proper gemmules with certain nascent cells, together with the superabundance of the gemmules derived from both parents, and the subsequent self-multiplication of the gemmules, 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 transposed or multiplied. For instance, a curious case has been recorded by Dr. Elliott Coues (27/58. 'Proc. Boston Soc. of Nat. Hist.' republished in 'Scientific Opinion' November 10, 1869 page 488.) of a monstrous chicken with a perfect additional RIGHT leg articulated to the LEFT side of the pelvis. Gold-fish often have supernumerary fins placed on various parts of their bodies. When the tail of a lizard is broken off, a double tail is sometimes reproduced; and when the foot of the salamander was divided longitudinally by Bonnet, additional digits were occasionally formed. Valentin injured the caudal extremity of an embryo, and three days afterwards it produced rudiments of a double pelvis and of double hind-limbs. (27/59. Todd 'Cyclop. of Anat. and Phys.' volume 4 1849-52 page 975.) When frogs, toads, etc., are born with their limbs doubled, as sometimes happens, the doubling, as Gervais remarks (27/60. 'Compte Rendus' November 14, 1865 page 800.), cannot be due to the complete fusion of two embryos, with the exception of the limbs, for the larvae are limbless. The same argument is applicable (27/61. As previously remarked by Quatrefages in his 'Metamorphoses de l'Homme' etc. 1862 page 129.) to certain insects produced with multiple legs or antennae, for these are metamorphosed from apodal or antennae-less larvae. Alphonse Milne-Edwards (27/62. Gunther 'Zoological Record' 1864 page 279.) has described the curious case of a crustacean in which one eye-peduncle supported, instead of a complete eye, only an imperfect cornea, and out of the centre of this a portion of an antenna was developed. A case has been recorded (27/63. Sedgwick 'Medico-Chirurg. Review' April 1863 page 454.) 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 (27/64. Isid. Geoffroy Saint-Hilaire 'Hist. des Anomalies' tome 1 1832 pages 435, 657; and tome 2 page 560.) of additional teeth having been developed in the orbit of the eye, and, more especially with horses, in the palate. Hairs occasionally appear in strange situations, as "within the substance of the brain." (27/65. Virchow 'Cellular Pathology' 1860 page 66.) Certain breeds of sheep bear a whole crowd of horns on their foreheads. As many as five spurs have been seen on both legs of certain Game-fowls. In the Polish fowl the male is ornamented with a topknot of hackles like those on his neck, whilst the female has a top-knot formed 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. Imperfect nails sometimes appear on the stumps of the amputated fingers of man (27/66. Muller 'Phys.' English Translation volume 1 1833 page 407. A case of this kind has lately been communicated to me.) and it is an interesting fact that with the snake-like Saurians, which present a series with more and more imperfect limbs, the terminations of the phalanges first disappear, "the nails becoming transferred to their proximal remnants, or even to parts which are not phalanges." (27/67. Dr. Furbringer 'Die Knochen etc. bei den schlangenahnlichen Sauriern' as reviewed in 'Journal of Anat. and Phys.' May 1870 page 286.)

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 of Vicia sativa replaced by 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 into petals, more or less completely, 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, pollen; for instance, I have seen the pollen-mass of an Ophrys, which is a very complex structure, developed in the edge 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. Mr. Salter and Dr. Maxwell Masters have found pollen within the ovules of the passion-flower and of the rose. Buds may be developed in the most unnatural positions, as on the petal of a flower. Numerous analogous facts could be given. (27/68. Moquin-Tandon 'Teratologie Veg.' 1841 pages 218, 220, 353. For the case of the pea see 'Gardener's Chronicle' 1866 page 897. With respect to pollen within ovules see Dr. Masters in 'Science Review' October 1873 page 369. The Rev. J.M. Berkeley describes a bud developed on a petal of a Clarkia in 'Gardener's Chronicle' April 28, 1866.)

I do not know how physiologists look at such facts as the foregoing. According to the doctrine of pangenesis, the gemmules of the transposed organs become 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 their elective affinities. Nor ought we to feel much surprise at the affinities of cells and gemmules varying, when we remember the many curious cases given in the seventeenth chapter, of plants which absolutely refuse to be fertilised by their own pollen, though abundantly fertile with that of any other individual of the same species, and in some cases only with that of a distinct species. It is manifest that the sexual elective affinities of such plants — to use the term employed by Gartner — have been modified. As the cells of adjoining or homologous parts will have nearly the same nature, they will be particularly liable to acquire by variation each other's elective affinities; and we can thus understand to a certain extent such cases as a crowd of horns on the heads of certain sheep, of several spurs on the legs of fowls, hackle-like feathers on the heads of the males of other fowls, and with the pigeon wing-like feathers on their legs and 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 other gemmules in due succession, as in the regrowth of an amputated limb. When parts which are homologous and similar in structure, as the vertebrae of snakes or the stamens of polyandrous flowers, etc., 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 Saint-Hilaire's law, that parts, which are already multiple, are extremely liable to vary in number.

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 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 plants, if their own pollen be excluded, are easily hybridised, so, in the case of cells, if the properly succeeding gemmules were absent, they would probably combine easily with other and allied gemmules, as we have just seen with transposed parts.

In variations caused by the direct action of changed conditions, of which several instances have been given, certain parts of the body are directly affected by the new conditions, and consequently throw off modified gemmules, which are transmitted to the offspring. On any ordinary view it is unintelligible how changed conditions, whether acting on the embryo, the young or the adult, 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 a 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 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 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 of the relation between the brain and the power of thought that we do not know positively whether a fixed habit induces any change in the nervous system, though this seems highly probable; but when such habit or other mental attribute, or insanity, is inherited, we must believe that some actual modification is transmitted (27/69. See some remarks to this effect by Sir H. Holland in his 'Medical Notes' 1839 page 32.); and this implies, according to our hypothesis, that gemmules derived from modified nerve-cells are transmitted to the offspring.

It is generally necessary that an organism should be exposed during several generations to changed conditions or habits, in order that any modification thus acquired should appear in the offspring. This may be partly due to the changes not being at first marked enough to catch 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 unmodified unit or part are transmitted in large numbers to successive generations, and that the gemmules derived from the same unit after it has been modified go on multiplying under the same favourable conditions which first caused the modification, until at last they become sufficiently numerous to overpower and supplant the old gemmules.

A difficulty may be here noticed; we have seen that there 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 plants propagated asexually should be far less variable than those propagated sexually. With respect to the direct action of changed conditions, we know that organisms produced from buds do not pass through the earlier phases of development; they will therefore not be exposed, at that period of life when structure is most readily modified, to the various causes inducing variability in the same manner as are embryos and young larval forms; but whether this is a sufficient explanation I know not.

With respect to variations due to reversion, there is a similar difference between plants propagated from buds and seeds. Many varieties can be propagated securely by buds, but generally or invariably revert to their parent-forms 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 these facts. Plants with variegated leaves, phloxes with striped flowers, barberries with seedless fruit, can all be securely propagated by buds taken from the stem or branches; but buds from the roots of these plants almost invariably lose their character and revert to their former condition. This latter fact is also inexplicable, unless buds developed from the roots are as distinct from those on the stem, as is one bud on the stem from another, and we know that these latter behave like independent organisms.