Professor Geddes's Theory of Variation in Plants.
In a paper read before the Edinburgh Botanical Society in 1886 Mr. Patrick Geddes laid down the outlines of a fundamental theory of plant variation, which he has further extended in the article "Variation and Selection" in the Encydopaedia Britannica, and in a paper read before the Linnaean Society but not yet published.
A theory of variation should deal alike with the origin of specific distinctions and with those vaster differences which characterise the larger groups, and he thinks it should answer such questions as—How an axis comes to be arrested to form a flower? how the various forms of inflorescence were evolved? how did perigynous or epigynous flowers arise from hypogynous flowers? and many others equally fundamental. Natural selection acting upon numerous accidental variations will not, he urges, account for such general facts as these, which must depend on some constant law of variation. This law he believes to be the well-known antagonism of vegetative and reproductive growth acting throughout the whole course of plant development; and he uses it to explain many of the most characteristic features of the structure of flowers and fruits.
Commencing with the origin of the flower, which all botanists agree in regarding as a shortened branch, he explains this shortening as an inevitable physiological fact, since the cost of the development of the reproductive elements is so great as necessarily to check vegetative growth. In the same manner the shortening of the inflorescence from raceme to spike or umbel, and thence to the capitulum or dense flower-head of the composite plants is brought about. This shortening, carried still further, produces the flattened leaf-like receptacle of Dorstenia, and further still the deeply hollowed fruity receptacle of the fig.
The flower itself undergoes a parallel modification due to a similar cause. It is formed by a series of modified leaves arranged round a shortened axis. In its earlier stages the number of these modified leaves is indefinite, as in many Ranunculaceae; and the axis itself is not greatly shortened, as in Myosurus. The first advance is to a definite number of parts and a permanently shortened axis, in the arrangement termed hypogynous, in which all the whorls are quite distinct from each other. In the next stage there is a further shortening of the central axis, leaving the outer portion as a ring on which the petals are inserted, producing the arrangement termed perigynous. A still further advance is made by the contraction of the axis, so as to leave the central part forming the ovary quite below the flower, which is then termed epigynous.
These several modifications are said to be parallel and definite, and to be determined by the continuous checking of vegetation by reproduction along what is an absolute groove of progressive change. This being the case, the importance of natural selection is greatly diminished. Instead of selecting and accumulating spontaneous indefinite variations, its function is to retard them after the stage of maximum utility has been independently reached. The same simple conception is said to unlock innumerable problems of vegetable morphology, large and small alike. It explains the inevitable development of gymnosperm into angiosperm by the checked vegetative growth of the ovule-bearing leaf or carpel; while such minor adaptations as the splitting fruit of the geranium or the cupped stigma of the pansy, can be no longer looked upon as achievements of natural selection, but must be regarded as naturally traceable to the vegetative checking of their respective types of leaf organ. Again, a detailed examination of spiny plants practically excludes the hypothesis of mammalian selection altogether, and shows spines to arise as an expression of the diminishing vegetativeness—in fact, the ebbing vitality of a species.[209]
Objections to the Theory.
The theory here sketched out is enticing, and at first sight seems calculated to throw much light on the history of plant development; but on further consideration, it seems wanting in definiteness, while it is beset with difficulties at every step. Take first the shortening of the raceme into the umbel and the capitulum, said to be caused by arrest of vegetative growth, due to the antagonism of reproduction. If this were the whole explanation of the phenomenon, we should expect the quantity of seed to increase as this vegetative growth diminished, since the seed is the product of the reproductive energy of the plant, and its quantity the best measure of that energy. But is this the case? The ranunculus has comparatively few seeds, and the flowers are not numerous; while in the same order the larkspur and the columbine have far more seeds as well as more flowers, but there is no shortening of the raceme or diminution of the foliage, although the flowers are large and complex. So, the extremely shortened and compressed flower-heads of the compositae produce comparatively few seeds—one only to each flower; while the foxglove, with its long spike of showy flowers, produces an enormous number.
Again, if the shortening of the central axis in the successive stages of hypogynous, perigynous, and epigynous flowers were an indication of preponderant reproduction and diminished vegetation, we should find everywhere some clear indications of this fact. The plants with hypogynous flowers should, as a rule, have less seed and more vigorous and abundant foliage than those at the other extreme with epigynous flowers. But the hypogynous poppies, pinks, and St. John's worts have abundance of seed and rather scanty foliage; while the epigynous dogwoods and honeysuckles have few seeds and abundant foliage. If, instead of the number of the seeds, we take the size of the fruit as an indication of reproductive energy, we find this at a maximum in the gourd family, yet their rapid and luxuriant growth shows no diminution of vegetative power. So that the statement that plant modifications proceed "along an absolute groove of progressive change" is contradicted by innumerable facts indicating advance and regression, improvement or degradation, according as the ever-changing environment renders one form more advantageous than the other. As one instance I may mention the Anonaceae or custard-apple tribe, which are certainly an advance from the Ranunculaceae; yet in the genus Polyalthea the fruit consists of a number of separate carpels, each borne on a long stalk, as if reverting to the primitive stalked carpellary leaves.
On the Origin of Spines.
But perhaps the most extraordinary application of the theory is that which considers spines to be an indication of the "ebbing vitality of a species," and which excludes "mammalian selection altogether." If this were true, spines should occur mainly in feeble, rare, and dying-out species, instead of which we have the hawthorn, one of our most vigorous shrubs or trees, with abundant vitality and an extensive range over the whole Palaearctic region, showing that it is really a dominant species. In North America the numerous thorny species of Crataegus are equally vigorous, as are the false acacia (Robinia) and the honey-locust (Gleditschia). Neither have the numerous species of very spiny Acacias been noticed to be rarer or less vigorous than the unarmed kinds.
On the other point—that spines are not due to mammalian selection—we are able to adduce what must be considered direct and conclusive evidence. For if spines, admittedly produced by aborted branches, petioles, or peduncles, are due solely or mainly to diminished vegetativeness or ebbing vitality, they ought to occur in all countries alike, or at all events in all whose similar conditions tend to check vegetation; whereas, if they are, solely or mainly, developed as a protection against the attacks of herbivorous mammals, they ought to be most abundant where these are plentiful, and rare or absent where indigenous mammalia are wanting. Oceanic islands, as compared with continents, would thus furnish a crucial test of the two theories; and Mr. Hemsley of Kew, who has specially studied insular floras, has given me some valuable information on this point. He says: "There are no spiny or prickly plants in the indigenous element of the St. Helena flora. The relatively rich flora of the Sandwich Isles is not absolutely without a prickly plant, but almost so. All the endemic genera are unarmed, and the endemic species of almost every other genus. Even such genera as Zanthoxylon, Acacia, Xylosoma, Lycium, and Solanum, of which there are many armed species in other countries, are only represented by unarmed species. The two endemic Rubi have the prickles reduced to the setaceous condition, and the two palms are unarmed.
"The flora of the Galapagos includes a number of prickly plants, among them several cacti (these have not been investigated and may be American species), but I do not think one of the known endemic species of any family is prickly or spiny.
"Spiny and prickly plants are also rare in New Zealand, but there are the formidably armed species of wild Spaniard (Aciphylla), one species of Rubus, the pungent-leaved Epacrideae and a few others."
Mr. J.G. Baker of Kew, who has specially studied the flora of Mauritius and the adjacent islands, also writes me on this point. He says: "Taking Mauritius alone, I do not call to mind a single species that is a spinose endemic tree or shrub. If you take the whole group of islands (Mauritius, Bourbon, Seychelles, and Rodriguez), there will be about a dozen species, but then nine of these are palms. Leaving out palms, the trees and shrubs of that part of the world are exceptionally non-spinose."
These are certainly remarkable facts, and quite inexplicable on the theory of spines being caused solely by checked vegetative growth, due to weakness of constitution or to an arid soil and climate. For the Galapagos and many parts of the Sandwich Islands are very arid, as is a considerable part of the North Island of New Zealand. Yet in our own moist climate and with our very limited number of trees and shrubs we have about eighteen spiny or prickly species, more, apparently, than in the whole endemic floras of the Mauritius, Sandwich Islands, and Galapagos, though these are all especially rich in shrubby and arboreal species. In New Zealand the prickly Rubus is a leafless trailing plant, and its prickles are probably a protection against the large snails of the country, several of which have shells from two to three and a half inches long.[210] The "wild Spaniards" are very spiny herbaceous Umbelliferae, and may have gained their spines to preserve them from being trodden down or eaten by the Moas, which, for countless ages, took the place of mammals in New Zealand. The exact use or meaning of the spines in palms is more doubtful, though they are, no doubt, protective against some animals; but it is certainly an extraordinary fact that in the entire flora of the Mauritius, so largely consisting of trees and shrubs, not a single endemic species should be thorny or spiny.
If now we consider that every continental flora produces a considerable proportion of spiny and thorny species, and that these rise to a maximum in South Africa, where herbivorous mammalia were (before the settlement of the country), perhaps, more abundant and varied than in any other part of the world; while another district, remarkable for well-armed vegetation, is Chile, where the camel-like vicugnas, llamas, and alpacas, and an abundance of large rodents wage perpetual war against shrubby vegetation, we shall see the full significance of the almost total absence of thorny and spiny plants in the chief oceanic islands; and so far from "excluding the hypothesis of mammalian selection altogether," we shall find in this hypothesis the only satisfactory explanation of the facts.
From the brief consideration of Professor Geddes's theory now given, we conclude that, although the antagonism between vegetative and reproductive growth is a real agency, and must be taken account of in our endeavour to explain many of the fundamental facts in the structure and form of plants, yet it is so overpowered and directed at every step by the natural selection of favourable variations, that the results of its exclusive and unmodified action are nowhere to be found in nature. It may be allowed to rank as one of those "laws of growth," of which so many have now been indicated, and which were always recognised by Darwin as underlying all variation; but unless we bear in mind that its action must always be subordinated to natural selection, and that it is continually checked, or diverted, or even reversed by the necessity of adaptation to the environment, we shall be liable to fall into such glaring errors as the imputing to "ebbing vitality" alone such a widespread phenomenon as the occurrence of spines and thorns, while ignoring altogether the influence of the organic environment in their production.[211]
The sketch now given of the chief attempts that have been made to prove that either the direct action of the environment or certain fundamental laws of variation are independent causes of modification of species, shows us that their authors have, in every case, failed to establish their contention. Any direct action of the environment, or any characters acquired by use or disuse, can have no effect whatever upon the race unless they are inherited; and that they are inherited in any case, except when they directly affect the reproductive cells, has not been proved. On the other hand, as we shall presently show, there is much reason for believing that such acquired characters are in their nature non-heritable.
Variation and Selection Overpower the Effects of Use and Disuse.
But there is another objection to this theory arising from the very nature of the effects produced. In each generation the effects of use or disuse, or of effort, will certainly be very small, while of this small effect it is not maintained that the whole will be always inherited by the next generation. How small the effect is we have no means of determining, except in the case of disuse, which Mr. Darwin investigated carefully. He found that in twelve fancy breeds of pigeons, which are often kept in aviaries, or if free fly but little, the sternum had been reduced by about one-seventh or one-eighth of its entire length, and that of the scapula about one-ninth. In domestic ducks the weight of the wing-bones in proportion to that of the whole skeleton had decreased about one-tenth. In domestic rabbits the bones of the legs were found to have increased in weight in due proportion to the increased weight of the body, but those of the hind legs were rather less in proportion to those of the fore legs than in the wild animal, a difference which may be imputed to their being less used in rapid motion. The pigeons, therefore, afford the greatest amount of reduction by disuse—one-seventh of the length of the sternum. But the pigeon has certainly been domesticated four or five thousand years; and if the reduction of the wings by disuse has only been going on for the last thousand years, the amount of reduction in each generation would be absolutely imperceptible, and quite within the limits of the reduction due to the absence of selection, as already explained. But, as we have seen in Chapter III, the fortuitous variation of every part or organ usually amounts to one-tenth, and often to one-sixth of the average dimensions—that is, the fortuitous variation in one generation among a limited number of the individuals of a species is as great as the cumulative effects of disuse in a thousand generations! If we assume that the effects of use or of effort in the individual are equal to the effects of disuse, or even ten or a hundred times greater, they will even then not equal, in each generation, the amount of the fortuitous variations of the same part. If it be urged that the effects of use would modify all the individuals of a species, while the fortuitous variations to the amount named only apply to a portion of them, it may be replied, that that portion is sufficiently large to afford ample materials for selection, since it often equals the numbers that can annually survive; while the recurrence in each successive generation of a like amount of variation would render possible such a rapid adjustment to new conditions that the effects of use or disuse would be as nothing in comparison. It follows, that even admitting the modifying effects of the environment, and that such modifications are inherited, they would yet be entirely swamped by the greater effects of fortuitous variation, and the far more rapid cumulative results of the selection of such variations.
Supposed Action of the Environment in Initiating Variations.
It is, however, urged that the reaction of the environment initiates variations, which without it would never arise; such, for instance, as the origin of horns through the pressures and irritations caused by butting, or otherwise using the head as a weapon or for defence. Admitting, for the sake of argument, that this is so, all the evidence we possess shows that, from the very first appearance of the rudiment of such an organ, it would vary to a greater extent than the amount of growth directly produced by use; and these variations would be subject to selection, and would thus modify the organ in ways which use alone would never bring about. We have seen that this has been the case with the branching antlers of the stag, which have been modified by selection, so as to become useful in other ways than as a mere weapon; and the same has almost certainly been the case with the variously curved and twisted horns of antelopes. In like manner, every conceivable rudiment would, from its first appearance, be subject to the law of variation and selection, to which, thenceforth, the direct effect of the environment would be altogether subordinate.
A very similar mode of reasoning will apply to the other branch of the subject—the initiation of structures and organs by the action of the fundamental laws of growth. Admitting that such laws have determined some of the main divisions of the animal and vegetable kingdom, have originated certain important organs, and have been the fundamental cause of certain lines of development, yet at every step of the process these laws must have acted in entire subordination to the law of natural selection. No modification thus initiated could have advanced a single step, unless it were, on the whole, a useful modification; while its entire future course would be necessarily subject to the laws of variation and selection, by which it would be sometimes checked, sometimes hastened on, sometimes diverted to one purpose, sometimes to another, according as the needs of the organism, under the special conditions of its existence, required such modification. We need not deny that such laws and influences may have acted in the manner suggested, but what we do deny is that they could possibly escape from the ever-present and all-powerful modifying effects of variation and natural selection.[212]
Weismann's Theory of Heredity.
Professor August Weismann has put forth a new theory of heredity founded upon the "continuity of the germ-plasm," one of the logical consequences of which is, that acquired characters of whatever kind are not transmitted from parent to offspring. As this is a matter of vital importance to the theory of natural selection, and as, if well founded, it strikes away the foundations of most of the theories discussed in the present chapter, a brief outline of Weismann's views must be attempted, although it is very difficult to make them intelligible to persons unfamiliar with the main facts of modern embryology.[213]
The problem is thus stated by Weismann: "How is it that in the case of all higher animals and plants a single cell is able to separate itself from amongst the millions of most various kinds of which an organism is composed, and by division and complicated differentiation to reconstruct a new individual with marvellous likeness, unchanged in many cases even throughout whole geological periods?" Darwin attempted to solve the problem by his theory of "Pangenesis," which supposed that every individual cell in the body gave off gemmules or germs capable of reproducing themselves, and that portions of these germs of each of the almost infinite number of cells permeate the whole body and become collected in the generative cells, and are thus able to reproduce the whole organism. This theory is felt to be so ponderously complex and difficult that it has met with no general acceptance among physiologists.
The fact that the germ-cells do reproduce with wonderful accuracy not only the general characters of the species, but many of the individual characteristics of the parents or more remote ancestors, and that this process is continued from generation to generation, can be accounted for, Weismann thinks, only on two suppositions which are physiologically possible. Either the substance of the parent germ-cell, after passing through a cycle of changes required for the construction of a new individual, possesses the capability of producing anew germ-cells identical with those from which that individual was developed, or the new germ-cells arise, as far as their essential and characteristic substance is concerned, not at all out of the body of the individual, but direct from the parent germ-cell. This latter view Weismann holds to be the correct one, and, on this theory, heredity depends on the fact that a substance of special molecular composition passes over from one generation to another. This is the "germ-plasm," the power of which to develop itself into a perfect organism depends on the extraordinary complication of its minutest structure. At every new birth a portion of the specific germ-plasm, which the parent egg-cell contains, is not used up in producing the offspring, but is reserved unchanged to produce the germ-cells of the following generation. Thus the germ-cells—so far as regards their essential part the germ-plasm—are not a product of the body itself, but are related to one another in the same way as are a series of generations of unicellular organisms derived from one another by a continuous course of simple division. Thus the question of heredity is reduced to one of growth. A minute portion of the very same germ-plasm from which, first the germ-cell, and then the whole organism of the parent, were developed, becomes the starting-point of the growth of the child.
The Cause of Variation.
But if this were all, the offspring would reproduce the parent exactly, in every detail of form and structure; and here we see the importance of sex, for each new germ grows out of the united germ-plasms of two parents, whence arises a mingling of their characters in the offspring. This occurs in each generation; hence every individual is a complex result reproducing in ever-varying degrees the diverse characteristics of his two parents, four grandparents, eight great-grandparents, and other more remote ancestors; and that ever-present individual variation arises which furnishes the material for natural selection to act upon. Diversity of sex becomes, therefore, of primary importance as the cause of variation. Where asexual generation prevails, the characteristics of the individual alone are reproduced, and there are thus no means of effecting the change of form or structure required by changed conditions of existence. Under such changed conditions a complex organism, if only asexually propagated, would become extinct. But when a complex organism is sexually propagated, there is an ever-present cause of change which, though slight in any one generation, is cumulative, and under the influence of selection is sufficient to keep up the harmony between the organism and its slowly changing environment.[214]
The Non-Heredity of Acquired Characters.
Certain observations on the embryology of the lower animals are held to afford direct proof of this theory of heredity, but they are too technical to be made clear to ordinary readers. A logical result of the theory is the impossibility of the transmission of acquired characters, since the molecular structure of the germ-plasm is already determined within the embryo; and Weismann holds that there are no facts which really prove that acquired characters can be inherited, although their inheritance has, by most writers, been considered so probable as hardly to stand in need of direct proof.
We have already shown, in the earlier part of this chapter, that many instances of change, imputed to the inheritance of acquired variations, are really cases of selection; while the very fact that use implies usefulness renders it almost impossible to eliminate the action of selection in a state of nature. As regards mutilations, it is generally admitted that they are not hereditary, and there is ample evidence on this point. When it was the fashion to dock horses' tails, it was not found that horses were born with short tails; nor are Chinese women born with distorted feet; nor are any of the numerous forms of racial mutilation in man, which have in some cases been carried on for hundreds of generations, inherited. Nevertheless, a few cases of apparent inheritance of mutilations have been recorded,[215] and these, if trustworthy, are difficulties in the way of the theory. The undoubted inheritance of disease is hardly a difficulty, because the predisposition to disease is a congenital, not an acquired character, and as such would be the subject of inheritance. The often-quoted case of a disease induced by mutilation being inherited (Brown-Sequard's epileptic guinea-pigs) has been discussed by Professor Weismann, and shown to be not conclusive. The mutilation itself—a section of certain nerves—was never inherited, but the resulting epilepsy, or a general state of weakness, deformity, or sores, was sometimes inherited. It is, however, possible that the mere injury introduced and encouraged the growth of certain microbes, which, spreading through the organism, sometimes reached the germ-cells, and thus transmitted a diseased condition to the offspring. Such a transference of microbes is believed to occur in syphilis and tuberculosis, and has been ascertained to occur in the case of the muscardine silkworm disease.[216]
The Theory of Instinct.
The theory now briefly outlined cannot be said to be proved, but it commends itself to many physiologists as being inherently probable, and as furnishing a good working hypothesis till displaced by a better. We cannot, therefore, accept any arguments against the agency of natural selection which are based upon the opposite and equally unproved theory that acquired characters are inherited; and as this applies to the whole school of what may be termed Neo-Lamarckians, their speculations cease to have any weight.
The same remark applies to the popular theory of instincts as being inherited habits; though Darwin gave very little weight to this, but derived almost all instincts from spontaneous useful variations which, like other spontaneous variations, are of course inherited. At first sight it appears as if the acquired habits of our trained dogs—pointers, retrievers, etc.—are certainly inherited; but this need not be the case, because there must be some structural or psychical peculiarities, such as modifications in the attachments of muscles, increased delicacy of smell or sight, or peculiar likes and dislikes, which are inherited; and from these, peculiar habits follow as a natural consequence, or are easily acquired. Now, as selection has been constantly at work in improving all our domestic animals, we have unconsciously modified the structure, while preserving only those animals which best served our purpose in their peculiar faculties, instincts, or habits.
Much of the mystery of instinct arises from the persistent refusal to recognise the agency of imitation, memory, observation, and reason as often forming part of it. Yet there is ample evidence that such agency must be taken into account. Both Wilson and Leroy state that young birds build inferior nests to old ones, and the latter author observes that the best nests are made by birds whose young remain longest in the nest. So, migration is now well ascertained to be effected by means of vision, long flights being made on bright moonlight nights when the birds fly very high, while on cloudy nights they fly low, and then often lose their way. Thousands annually fly out to sea and perish, showing that the instinct to migrate is imperfect, and is not a good substitute for reason and observation.
Again, much of the perfection of instinct is due to the extreme severity of the selection during its development, any failure involving destruction. The chick which cannot break the eggshell, the caterpillar that fails to suspend itself properly or to spin a safe cocoon, the bees that lose their way or that fail to store honey, inevitably perish. So the birds that fail to feed and protect their young, or the butterflies that lay their eggs on the wrong food-plant, leave no offspring, and the race with imperfect instincts perishes. Now, during the long and very slow course of development of each organism, this rigid selection at every step of progress has led to the preservation of every detail of structure, faculty, or habit that has been necessary for the preservation of the race, and has thus gradually built up the various instincts which seem so marvellous to us, but which can yet be shown to be in many cases still imperfect. Here, as everywhere else in nature, we find comparative, not absolute perfection, with every gradation from what is clearly due to imitation or reason up to what seems to us perfect instinct—that in which a complex action is performed without any previous experience or instruction.[217]
Having now passed in review the more important of the recent objections to, or criticisms of, the theory of natural selection, we have arrived at the conclusion that in no one case have the writers in question been able materially to diminish its importance, or to show that any of the laws or forces to which they appeal can act otherwise than in strict subordination to it. The direct action of the environment as set forth by Mr. Herbert Spencer, Dr. Cope, and Dr. Karl Semper, even if we admit that its effects on the individual are transmitted by inheritance, are so small in comparison with the amount of spontaneous variation of every part of the organism that they must be quite overshadowed by the latter. And if such direct action may, in some cases, have initiated certain organs or outgrowths, these must from their very first beginnings have been subject to variation and natural selection, and their further development have been almost wholly due to these ever-present and powerful causes. The same remark applies to the views of Professor Geddes on the laws of growth which have determined certain essential features in the morphology of plants and animals. The attempt to substitute these laws for those of variation and natural selection has failed in cases where we can apply a definite test, as in that of the origin of spines on trees and shrubs; while the extreme diversity of vegetable structure and form among the plants of the same country and of the same natural order, of itself affords a proof of the preponderating influence of variation and natural selection in keeping the many diverse forms in harmony with the highly complex and ever-changing environment.
Lastly, we have seen that Professor Weismann's theory of the continuity of the germ-plasm and the consequent non-heredity of acquired characters, while in perfect harmony with all the well-ascertained facts of heredity and development, adds greatly to the importance of natural selection as the one invariable and ever-present factor in all organic change, and that which can alone have produced the temporary fixity combined with the secular modification of species. While admitting, as Darwin always admitted, the co-operation of the fundamental laws of growth and variation, of correlation and heredity, in determining the direction of lines of variation or in the initiation of peculiar organs, we find that variation and natural selection are ever-present agencies, which take possession, as it were, of every minute change originated by these fundamental causes, check or favour their further development, or modify them in countless varied ways according to the varying needs of the organism. Whatever other causes have been at work, Natural Selection is supreme, to an extent which even Darwin himself hesitated to claim for it. The more we study it the more we are convinced of its overpowering importance, and the more confidently we claim, in Darwin's own words, that it "has been the most important, but not the exclusive, means of modification."
[198] See the Duke of Argyll's letter in Nature, vol. xxxiv. p. 336.
[199] Journal of the Anthropological Institute, vol. xv. pp. 246-260.
[200] The idea of the non-heredity of acquired variations was suggested by the summary of Professor Weismann's views, in Nature, referred to later on. But since this chapter was written I have, through the kindness of Mr. E.B. Poulton, seen some of the proofs of the forthcoming translation of Weismann's Essays on Heredity, in which he sets forth an explanation very similar to that here given. On the difficult question of the almost entire disappearance of organs, as in the limbs of snakes and of some lizards, he adduces "a certain form of correlation, which Roux calls 'the struggle of the parts in the organism,'" as playing an important part. Atrophy following disuse is nearly always attended by the corresponding increase of other organs: blind animals possess more developed organs of touch, hearing, and smell; the loss of power in the wings is accompanied by increased strength of the legs, etc. Now as these latter characters, being useful, will be selected, it is easy to understand that a congenital increase of these will be accompanied by a corresponding congenital diminution of the unused organ; and in cases where the means of nutrition are deficient, every diminution of these useless parts will be a gain to the whole organism, and thus their complete disappearance will, in some cases, be brought about directly by natural selection. This corresponds with what we know of these rudimentary organs.
It must, however, be pointed out that the non-heredity of acquired characters was maintained by Mr. Francis Galton more than twelve years ago, on theoretical considerations almost identical with those urged by Professor Weismann; while the insufficiency of the evidence for their hereditary transmission was shown, by similar arguments to those used above and in the work of Professor Weismann already referred to (see "A Theory of Heredity," in Journ. Anthrop. Instit., vol. v. pp. 343-345).
[201] This explanation is derived from Weismann's Theory of the Continuity of the Germ-Plasm as summarised in Nature.
[202] See a collection of his essays under the title, The Origin of the Fittest: Essays on Evolution, D. Appleton and Co. New York. 1887.
[203] Origin of the Fittest, p. 174.
[204] Ibid. p. 29. It may be here noted that Darwin found these theories unintelligible. In a letter to Professor E.T. Morse in 1877, he writes: "There is one point which I regret you did not make clear in your Address, namely, what is the meaning and importance of Professors Cope and Hyatt's views on acceleration and retardation? I have endeavoured, and given up in despair, the attempt to grasp their meaning" (Life and Letters, vol. iii. p. 233).
[205] Origin of the Fittest, p. 374.
[206] Origin of the Fittest, p. 40.
[207] The Natural Conditions of Existence as they Affect Animal Life. London, 1883.
[208] In Dr. Weismann's essay on "Heredity," already referred to, he considers it not improbable that changes in organisms produced by climatic influences may be inherited, because, as these changes do not affect the external parts of an organism only, but often, as in the case of warmth or moisture permeate the whole structure, they may possibly modify the germ-plasm itself, and thus induce variations in the next generation. In this way, he thinks, may possibly be explained the climatic varieties of certain butterflies, and some other changes which seem to be effected by change of climate in a few generations.
[209] This brief indication of Professor Geddes's views is taken from the article "Variation and Selection" in the Encyclopedia Britannica, and a paper "On the Nature and Causes of Variation in Plants" in Trans. and Proc. of the Edinburgh Botanical Society, 1886; and is, for the most part, expressed in his own words.
[210] Placostylis bovinus, 3½ inches long; Paryphanta Busbyi, 3 in. diam.; P. Hochstetteri, 2¾ in. diam.
[211] The general arguments and objections here set forth will apply with equal force to Professor G. Henslow's theory of the origin of the various forms and structures of flowers as due to "the responsive actions of the protoplasm in consequence of the irritations set up by the weights, pressures, thrusts, tensions, etc., of the insect visitors" (The Origin of Floral Structures through Insect and other Agencies, p. 340). On the assumption that acquired characters are inherited, such irritations may have had something to do with the initiation of variations and with the production of certain details of structure, but they are clearly incompetent to have brought about the more important structural and functional modifications of flowers. Such are, the various adjustments of length and position of the stamens to bring the pollen to the insect and from the insect to the stigma; the various motions of stamens and styles at the right time and the right direction; the physiological adjustments bringing about fertility or sterility in heterostyled plants; the traps, springs, and complex movements of various parts of orchids; and innumerable other remarkable phenomena.
For the explanation of these we have no resource but variation and selection, to the effects of which, acting alternately with regression or degradation as above explained (p. 328) must be imputed the development of the countless floral structures we now behold. Even the primitive flowers, whose initiation may, perhaps, have been caused, or rendered possible, by the irritation set up by insects' visits, must, from their very origin, have been modified, in accordance with the supreme law of utility, by means of variation and survival of the fittest.
[212] In an essay on "The Duration of Life," forming part of the translation of Dr. Weismann's papers already referred to, the author still further extends the sphere of natural selection by showing that the average duration of life in each species has been determined by it. A certain length of life is essential in order that the species may produce offspring sufficient to ensure its continuance under the most unfavourable conditions; and it is shown that the remarkable inequalities of longevity in different species and groups may be thus accounted for. Yet more, the occurrence of death in the higher organisms, in place of the continued survival of the unicellular organisms however much they may increase by subdivision, may be traced to the same great law of utility for the race and survival of the fittest. The whole essay is of exceeding interest, and will repay a careful perusal. A similar idea occurred to the present writer about twenty years back, and was briefly noted down at the time, but subsequently forgotten.
[213] The outline here given is derived from two articles in Nature, vol. xxxiii. p. 154, and vol. xxxiv. p. 629, in which Weismann's papers are summarised and partly translated.
[214] There are many indications that this explanation of the cause of variation is the true one. Mr. E.B. Poulton suggests one, in the fact that parthenogenetic reproduction only occurs in isolated species, not in groups of related species; as this shows that parthenogenesis cannot lead to the evolution of new forms. Again, in parthenogenetic females the complete apparatus for fertilisation remains unreduced; but if these varied as do sexually produced animals, the organs referred to, being unused, would become rudimentary.
Even more important is the significance of the "polar bodies," as explained by Weismann in one of his Essays; since, if his interpretation of them be correct, variability is a necessary consequence of sexual generation.
[215] Darwin's Animals and Plants, vol. ii. pp. 23, 24.
[216] In his essay on "Heredity," Dr. Weismann discusses many other cases of supposed inheritance of acquired characters, and shows that they can all be explained in other ways. Shortsightedness among civilised nations, for example, is due partly to the absence of selection and consequent regression towards a mean, and partly to its individual production by constant reading.
[217] Weismann explains instinct on similar lines, and gives many interesting illustrations (see Essays on Heredity). He holds "that all instinct is entirely due to the operation of natural selection, and has its foundation, not upon inherited experiences, but upon variations of the germ." Many interesting and difficult cases of instinct are discussed by Darwin in Chapter VIII of the Origin of Species, which should be read in connection with the above remarks.
Since this chapter was written my attention has been directed to Mr. Francis Galton's Theory of Heredity (already referred to at p. 417) which was published thirteen years ago as an alternative for Darwin's theory of pangenesis.
Mr. Galton's theory, although it attracted little attention, appears to me to be substantially the same as that of Professor Weismann. Galton's "stirp" is Weismann's "germ-plasm." Galton supposes the sexual elements in the offspring to be directly formed from the residue of the stirp not used up in the development of the body of the parent—Weismann's "continuity of the germ-plasm." Galton also draws many of the same conclusions from his theory. He maintains that characters acquired by the individual as the result of external influences cannot be inherited, unless such influences act directly on the reproductive elements—instancing the possible heredity of alcoholism, because the alcohol permeates the tissues and may reach the sexual elements. He discusses the supposed heredity of effects produced by use or disuse, and explains them much in the same manner as does Weismann. Galton is an anthropologist, and applies the theory, mainly, to explain the peculiarities of hereditary transmission in man, many of which peculiarities he discusses and elucidates. Weismann is a biologist, and is mostly concerned with the application of the theory to explain variation and instinct, and to the further development of the theory of evolution. He has worked it out more thoroughly, and has adduced embryological evidence in its support; but the views of both writers are substantially the same, and their theories were arrived at quite independently. The names of Galton and Weismann should therefore be associated as discoverers of what may be considered (if finally established) the most important contribution to the evolution theory since the appearance of the Origin of Species.
DARWINISM APPLIED TO MAN