by Grant Allen
A year later, in 1876, came the ‘Effects of Cross and Self Fertilisation in the Vegetable Kingdom.’ So far as regarded the world of plants, especially with respect to its higher divisions, this work was of immense theoretical importance; and it also cast a wonderful side-light upon the nature of that strange distinction of sex which occurs both in the vegetable and animal kingdom, and in each is the concomitant — one might almost say the necessary concomitant — of high development and complex organisation. The great result attained by Darwin in his long and toilsome series of experiments on this interesting subject was the splendid proof of the law that cross-fertilisation produces finer and healthier offspring, while continuous self-fertilisation tends in the long run to degradation, degeneration, and final extinction.
Here as elsewhere, however, Darwin’s principle does not spring spontaneous, like Athene from the head of Zeus, a goddess full-formed, uncaused, inexplicable: it arises gradually by a slow process of development and modification from the previous investigations of earlier biologists. At the close of the last century, in the terrible year of upheaval 1793, a quiet German botanist, Christian Konrad Sprengel by name, published at Berlin his long unheeded but intensely interesting work on the ‘Fertilisation of Flowers.’ In the summer of 1789, while all Europe was ablaze with the news that the Bastille had been stormed, and a new era of humanity begun, the calm and peaceful Pomeranian observer was noting in his own garden the curious fact that many flowers are incapable of being fertilised without the assistance of flying insects, which carry pollen from the stamens of one blossom to the sensitive surface or ovary of the next. Hence he concluded that the secretion of honey or nectar in flowers, the contrivances by which it is protected from rain, the bright hues or lines of the corolla, and the sweet perfume distilled by the blossoms, are all so many cunning devices of nature to ensure fertilisation by the insect-visitors. Moreover, Sprengel observed that many flowers are of one sex only, and that in several others the sexes do not mature simultaneously; ‘so that,’ said he, ‘nature seems to intend that no flower shall be fertilised by means of its own pollen.’ Indeed, in some instances, as he showed by experiments upon the yellow day lily, plants impregnated from their own stamens cannot be made to set seed at all. ‘So near,’ says his able successor, Hermann Müller, ‘was Sprengel to the distinct recognition of the fact that self-fertilisation leads to worse results than cross-fertilisation, and that all the arrangements which favour insect-visits are of value to the plant itself, simply because the insect-visitors effect cross-fertilisation!’ As in most other anticipatory cases, however, it must be here remarked that Sprengel’s idea was wholly teleological: he conceived of nature as animated by a direct informing principle, which deliberately aimed at a particular result; whereas Darwin rather came to the conclusion that cross-fertilisation as a matter of fact does actually produce beneficial results, and that therefore those plants which varied most in the direction of arrangements for favouring insect-visits were likely to be exceptionally fortunate in the struggle for existence against competitors otherwise arranged. It is just the usual Darwinian substitution of an efficient for a final cause.
Even before Sprengel, Kölreuter had recognised, in 1761, that self-fertilisation was avoided in nature; and his observations and experiments on intercrossing and on hybridism were largely relied upon by Darwin himself, to whom they suggested at an early period many fruitful lines of original investigation. In 1799, again, Andrew Knight, following up the same line of thought in England as Sprengel in Germany, declared as the result of his close experiments upon the garden pea, that no plant ever fertilises itself for a perpetuity of generations. But Knight’s law, not being brought into causal connection with any great fundamental principle of nature, was almost entirely overlooked by the scientific world until the publication of Darwin’s ‘Origin of Species,’ half a century later. The same neglect also overtook Sprengel’s immensely interesting and curious work on fertilisation of flowers. The world, in fact, was not yet ready for the separate treatment of functional problems connected with the interrelations of organic beings; so Knight and Sprengel were laid aside unnoticed on the dusty top bookshelves of public libraries, while the dry classificatory and systematic biology of the moment had it ail its own way for the time being on the centre reading-tables. So many separate and independent strands of thought does it ultimately require to make up the grand final generalisation which the outer world attributes in its totality to the one supreme organising intelligence.
But in the ‘Origin of Species’ itself Darwin reiterated and emphasised Knight’s law as a general and all-pervading principle of nature, placing it at the same time on broader and surer biological foundations by affiliating it intimately upon his own great illuminating and unifying doctrine of natural selection. He also soon after rescued from oblivion Sprengel’s curious and fairy-like book, showing in full detail in his work on orchids the wonderful contrivances by which flowers seek to attract and to secure the assistance of insects for the impregnation of their embryo seeds. In the ‘Variation of Animals and Plants under Domestication,’ he further showed that breeding in-and-in diminishes the strength and productiveness of the offspring; while crossing with another stock produces, on the contrary, the best possible physical results in both directions. And now at last, in the ‘Effects of Cross and Self fertilisation,’ he proved by careful and frequently repeated experiments that a constant infusion of fresh blood (so to speak) is essential to the production of the healthiest offspring. In the words of his own emphatic summing up, ‘Nature abhors perpetual self-fertilisation.’
The immediate result of these new statements and this fresh rationale of Knight’s law was to bring down Sprengel forthwith from the top shelf, where he had languished ingloriously for seventy years, and to set a whole school of ardent botanical observers working hard in the lines he had laid down upon the mutual correlations of insects and flowers. A vast literature sprang up at once upon this enchanting and long-neglected subject, the most eminent workers in the rediscovered field being Delpino in Italy, Hildebrand and Hermann Müller in Germany, Axel in Sweden, Lubbock in England, and Fritz Müller in tropical South America. Darwin found the question, in fact, almost taken out of his hands before he had time himself to treat of it; for Hildebrand’s chief work was published as early as 1867, while Axel’s appeared in 1869, both of them several years earlier than Darwin’s own final essay on the subject in the ‘Effects of Cross and Self Fertilisation.’ No statement, perhaps, could more clearly mark the enormous impetus given to researches in this direction than the fact that D’Arcy Thompson, in his appendix to Müller’s splendid work on the ‘Fertilisation of Flowers,’ has collected a list of no less than eight hundred and fourteen separate works or important papers bearing on that special department of botany, almost all of them subsequent in date to the first publication of the ‘Origin of Species.’ So widely did the Darwinian wave extend, and so profoundly did it affect every minute point of biological and psychological investigation.
Each of these later works of Darwin’s consists, as a rule, of an expansion of some single chapter or paragraph in the ‘Origin of Species;’ or, to speak more correctly, of an arrangement of the materials collected and the experiments designed for that particular portion of the great projected encyclopædia of evolutionism, of which the ‘Origin of Species’ itself was but a brief anticipatory summary or rough outline. Thus, the book on Orchids, published in 1862, is already foreshadowed in a part of the chapter on the Difficulties of the Theory of Natural Selection; the ‘Movements and Habits of Climbing Plants’ (1865) is briefly summarised by anticipation in the long section on Modes of Transition; the ‘Variation of Animals and Plants under Domestication’ (1868) consists of the vast array of pièces justificatives for the first chapter of the ‘Origin of Species;’ and the germ of the ‘Cross and Self Fertilisation’ (1876) is to be seen in the passage ‘On the Intercrossing of Individuals’ in Chapter IV. of the same work. It was well indeed
that Darwin began by publishing the shorter and more manageable abstract; the half, as the wise Greek proverb shrewdly remarks, is often more than the whole; and a world that eagerly devoured the first great deliverance of the Darwinian principle, might have stood aghast had it been asked to swallow it piecemeal in such gigantic treatises as those with which its author afterwards sought thrice to vanquish all his foes and thrice to slay the slain.
Yet, with each fresh manifestation of Darwin’s inexhaustible resources, on the other hand, the opposition to his principles grew feebler and feebler, and the universality of their acceptance more and more pronounced, till at last, among biologists at least, not to be a Darwinian was equivalent to being hopelessly left behind by the general onward movement of the time. In 1874 Tyndall delivered his famous address at the Belfast meeting of the British Association; and in 1877, from the same presidential chair at Plymouth, Allen Thomson, long reputed a doubtful waverer, enforced his cordial adhesion to the Darwinian principles by his inaugural discourse on ‘The Development of the Forms of Animal Life.’ A new generation of active workers, trained up from the first in the evolutionary school, like Romanes, Ray Lankester, Thistleton Dyer, Balfour, Sully, and Moggridge, had now risen gradually around the great master; and in every direction he could see the seed he had himself planted being watered and nourished in fresh soil by a hundred ardent and enthusiastic young disciples. Even in France, ever irresponsive to the touch of new ideas of alien origin, Colonel Moulinié’s admirable and sympathetic translations were beginning to win over to the evolutionary creed many rising workers; while in Germany, Victor Carus’s excellent versions had from the very first brought in the enthusiastic Teutonic biologists with a congenial ‘swarmery’ to the camp of the Darwinians. Correspondents from every part of the world kept pressing fresh facts and fresh applications upon the founder of the faith; and Darwin saw his own work so fast being taken out of his hands by specialist disciples that he abandoned entirely his original intention of publishing in detail the basis of his first book, and contented himself instead with tracing out minutely some minor portions of his contemplated task as specimens of evolutionary method.
In 1877, in pursuance of this changed purpose, Darwin published his book on ‘Forms of Flowers,’ in which he dealt closely with the old problem of differently shaped blossoms on plants of the same species. It had long been known, to take a single example, that primroses existed in two forms, the pin-eyed and the thrum-eyed, of which the former has the pin-like summit of the pistil at the top of the tube, and the stamens concealed half way down its throat; while in the latter these relative positions are exactly reversed, the stamens answering in place to the pistil of the alternative form with geometrical accuracy. As early as 1862 Darwin had shown, in the ‘Journal of the Linnean Society,’ that this curious arrangement owed its development to the greater security which it afforded for cross-fertilisation, because in this way each flower had to be impregnated with the pollen, from a totally distinct blossom, growing on a different individual plant. In a series of successive papers read before the same Society in the years between 1863 and 1868, he had extended a similar course of explanation to the multiform flowers of the flaxes, the loosestrifes, the featherfoil, the auricula, the buckbean, and several other well-known plants. At last, in 1877, he gathered together into one of the now familiar green-covered volumes the whole of his observations on this strange peculiarity, and proved by abundant illustration and experiment that the diversity of form is always due through natural selection to the advantage gained by perfect security of cross-fertilisation, resulting as it invariably does in the production of the finest, strongest, and most successful seedlings. Any variation, however peculiar, which helps to ensure this constant infusion of fresh blood is certain to be favoured in the struggle for life, owing to the superior vitality of the stock it begets. But it is worthy of notice, as showing the extreme minuteness and exhaustiveness of Darwin’s method on the small scale, side by side with his extraordinary and unusual power of rising to the very highest and grandest generalisations, that the volume which he devoted to the elucidation of this minor factor in the question of hereditary advantages runs to nearly as many pages as the last edition of the ‘Origin of Species’ itself. So great was the wealth of observation and experiment which he could lavish upon the solution of a single, small, incidental problem.
Even fuller in minute original research was the work which Darwin published in 1880, on ‘The Power of Movement in Plants,’ detailing the result of innumerable observations on the seemingly irresponsible yet almost purposive rotations of the growing rootlets and young stems of peas and climbers. Anyone who wishes to see on what a wide foundation of irrefragable fact the great biologist built up the stately fabric of his vast theories cannot do better than turn for instruction to this remarkable volume, which the old naturalist gave to the world some time after passing the allotted span of threescore years and ten.
It was in the same year (1880) that Huxley delivered at the Royal Institution his famous address on the Coming of Age of the ‘Origin of Species.’ The time was a favourable one for reviewing the silent and almost unobserved progress of a great revolution. Twenty-one years had come and gone since the father of modern scientific evolutionism had launched upon the world his tentative work. In those twenty-one years the thought of humanity had been twisted around as upon some invisible pivot, and a new heaven and a new earth had been presented to the eyes of seers and thinkers. One-and-twenty years before, despite the influence of Hutton and of Lyell, the dominant view of the earth’s past history revealed but one vast and lawless succession of hideous catastrophes. Wholesale creations and wholesale extinctions, world-wide cataclysms followed by fresh world-wide births of interwoven faunas and floras — these, said Huxley, were the ordinary machinery of the geological epic brought into fashion by the misapplied genius of the mighty Cuvier. One-and-twenty years after, the opponents themselves had given up the game in its fullest form as lost beyond the hope of possible restitution. Some hesitating thinkers, it is true, while accepting the evolutionary doctrine more or less in its earlier form, like Mivart and Meehan, yet refused their assent on one ground or another to the specific Darwinian doctrine of natural selection. Others, like Wallace, made a special exception with regard to the development of the human species, which they supposed to be due to other causes from those implied in the remainder of the organic scale. Yet on the whole, biological science had fairly carried the day in favour of evolution, in one form or another, and not even the cavillers dared now to suggest that whole systems of creation had been swept away en bloc, and remade again in different forms for a succeeding epoch, in accordance with the belief which was almost universal among geologists up to the exact moment of the publication of Darwin’s masterpiece.
During the twenty-one years, too, as Huxley likewise pointed out, an immense number of new facts had come to strengthen the hands of the evolutionists at the very point where they had before felt themselves most openly vulnerable. Palæontology had supplied many of those missing links in the organic chain whose absence from the interrupted and imperfect geological record had been loudly alleged against the Darwinian hypothesis in the earlier days of struggle and hesitation. Two years after the publication of the ‘Origin of Species,’ the discovery of a winged and feathered creature, happily preserved for us in the Solenhofen slates, with lizard-like head and teeth and tail, and bird-like pinions, feet, and breast, had bridged over in part the great gap that yawns between the existing birds and reptiles. A few years later, new fossil reptilian forms, erect on their hind legs like kangaroos, and with very singular peculiarities of bony structure, had helped still further to show the nature of the modifications by which the scale-bearing quadruped type passed slowly into that of the feather-bearing biped. In 1875, again, Professor Marsh’s discovery of the toothed birds in the American cretaceous strata completed the illustrative series of transitional forms over what had once been the most remarkable existing bre
ak in the continuity of organic development. Similarly, Hofmeister’s investigations in the vegetable world brought close together the flowering and flowerless plants, by indicating that the ferns and the horsetails were connected in curious unforeseen ways, through the pill-worts and club-mosses, with the earliest and simplest of forest trees, the firs and the puzzle-monkeys. In minor matters like progress was continually reported on every side. Gaudry found among the fossils of Attica the successive stages by which the ancient and undeveloped civets passed into the more modern and specialised tribe of the hyænas; Marsh traced out in Western America the ancestry of the horse from a five-toed creature no bigger than a fox, through intermediate four-toed and three-toed forms, to the existing single solid-hoofed type with its digits reduced to the minimum of unity; and Filhol unearthed among the phosphorites of Quercy the common progenitor of the most distinct among the recent carnivores, the cats and the dogs, the plantigrade bears and the digitigrade pumas. ‘So far as the animal world is concerned,’ Professor Huxley said in conclusion, reviewing these additions to the evidence upon that memorable occasion, ‘evolution is no longer a speculation but a statement of historical fact.’ Of Darwin himself he remarked truly, ‘He has lived long enough to outlast detraction and opposition, and to see the stone that the builders rejected become the head-stone of the corner.’