1741-1789 Characteristics of the Period.The chief historical events are the decline of the French monarchy, the French revolution, the rise of Prussia, the expansion of England, and the American Declaration of Independence. In the history of thought we remark the introduction of the historical or comparative method, which seeks to co-ordinate facts and to trace events to their causes. Science steadily grows in influence, and freethought wins many triumphs; this is the age of Voltaire, Rousseau, and the EncyclopÆdists, of David Hume, of the French economists and Adam Smith. Systems of Flowering Plants: LinnÆus and the Jussieus.LinnÆus is remembered as a man of great industry, enterprise, and sagacity, who was inspired from boyhood by a passion for natural history and spent a long life in advancing it. He was early recognised as a leader in more than one branch of the study. L'Obel, Morison, and Ray had laboured to found a natural system of flowering plants, and it was they who laid the foundation upon which all their successors have built. The work did not, however, go steadily forward on the original plan. When LinnÆus entered upon the scene the prevalent systems were only moderately natural, and far from convenient in practice. To place the undescribed species which poured in from North America and other distant countries was a difficult task, LinnÆus gained currency for his system by connecting it with the newly accepted doctrine of sexes in plants. That doctrine was not conceived nor demonstrated by him (see p. 48), and it had, as we now see, no further connection with classification by stamens and styles than that it explained the almost universal occurrence of such parts in flowering plants. But LinnÆus had persuaded himself that he had done more to establish the existence of sexes in plants than anybody else, and that the physiological importance of stamens and styles was a proof of their systematic value. Neither of these beliefs can stand inquiry, but both were extremely influential on contemporary opinion. The so-called Sexual System achieved an immense success everywhere but in France and Germany. Botanists of small experience were now able to say whether the plants which seemed to be new were really undescribed or not; if undescribed, what was their appropriate place in the system. The congestion of systematic botany was relieved. The great naturalist appealed to posterity by publishing the sketch of a natural system of flowering plants, which he accompanied by judicious expositions of the philosophy of classification. He had the permanent reform of systematic botany really at heart; he did not believe that his own Sexual System could be final; and he was glad to help in setting up a better one. To this end he united groups of genera into families which he did not pretend to define, being often guided only by an obscure sense of natural bonds of union. Bernard de Jussieu, one of the most patient and observant of systematists, devoted his life to the same task, and profited by the example of LinnÆus. He published nothing, but found expression for his views in the arrangement of a botanic garden at Versailles. His ideas were afterwards developed by his nephew, A. L. de Jussieu, in the Genera Plantarum (1789). Affinity became at length the avowed basis of every botanical system. No convenience in practice, no agreement or difference in habit, was knowingly permitted to override this mysterious property. What then is affinity? What are natural groups of animals and plants, and how do they arise? Until the year LinnÆus' imperfect appreciation of the fundamental difference between a natural classification of living things and such classifications as man makes for his own practical ends is further revealed by his admission of a third kingdom of nature. LinnÆus had a passionate love of order and clearness, enforced by an inexhaustible power of work. Hence he was able to serve his own generation with great effect, to methodise the labours of naturalists, to devise useful expedients for lightening their toil (such as his strict binomial nomenclature), From an engraving (1779) after the portrait by Roslin. RÉaumur and the History of Insects.RÉaumur was born to wealth, and made timely use of his leisure to study the sciences and win for himself a place among natural philosophers. His inclinations directed him first towards mathematics, physics, and, a little later, towards the practical arts. He took a leading part in a magnificent description of French industries, which had been undertaken by the AcadÉmie des Sciences. Not content with describing the processes in use, he perpetually laboured to improve them. The manufacture of steel, tin-plate, and porcelain, the hanging of carriages and the fitting of axles, the improvement of the thermometer, glass hives, and the hatching of fowls' eggs by artificial heat are among the many objects to which his attention was directed. Natural History gradually took a more and more prominent place in his studies, and a great History of Insects engaged the last years of his busy life. RÉaumur was neither an anatomist nor a systematist, at least he gained no distinction in either of these branches of biology. No biological laboratory had been dreamt of in his day; he lacked the manipulative skill of Swammerdam or Lyonet; he was no draughtsman, No abstract of RÉaumur's Histoire des Insectes is possible, but we may at least give one example of his mode of treatment. Let us select his account of the proboscis of a moth, the first full account that was ever Some moths have a tape-like proboscis; in others it is cylindrical. It can be made to protrude by gentle pressure on the head, or be unrolled by a pin passed into the centre of the spire; it is composed of innumerable joints, and tapers from the base to the tip. When forcibly unrolled, it often splits lengthwise into halves. At the time of escape from the chrysalis the halves are always free, and they require careful adjustment in order that a continuous sucking-tube may be obtained. A newly emerged moth may be seen to roll and unroll its proboscis repeatedly, until at last the halves cohere in the proper position. Sometimes they begin to dry before the operation is completed, the half-tubes get curled, and then the unfortunate moth becomes incapable of feeding at all. Each half is a demi-canal, whose meeting edges interlock by minute hooks. The mechanism reminds RÉaumur of that which connects RÉaumur was careful to correct his anatomical studies by close observation of the live insect. He reared an angle-shades moth, which he kept several days without food. When he saw it repeatedly extending its proboscis, he put near it a piece of sugar. The moth at once began to suck, and became so absorbed in satisfying its hunger that it allowed RÉaumur to carry it on a sheet of paper to a window and to examine it closely with a lens. The proboscis was sometimes extended for several minutes at a time, and then rolled up for an instant; its tip was either employed in exploring the surface or closely applied to the sugar. By means of the lens a slender column of liquid was seen to pass along the central canal towards the head. Now and then, however, a limpid fluid was seen to pass down the proboscis; this was the saliva which was used to moisten the sugar, and then sucked up again. The Budding-out of New Animals (Hydra): another Form of Propagation without Mating (Aphids).In the year 1744 a young Genevese, Abraham Trembley, tutor in the family of Bentinck, who was then English resident at the Hague, rose into sudden fame by a solid and well-timed contribution to natural history. Trembley and his pupils used to fish for aquatic insects in the ponds belonging to the residence, and in the summer of 1740 he happened to collect some water-weeds, which he put into a glass vessel and set in a window. When the floating objects had come to After four years of observation a handsome quarto volume was published, which told the history of "The freshwater Polyp," a name suggested by RÉaumur; the Latin name of Hydra was given by LinnÆus. Hydra had been discovered and slightly described forty years before by Leeuwenhoek, who had seen two young polyps branching from one parent and spontaneously becoming free. Trembley made out all that a simple lens, guided by a skilful hand and a keen eye, could discover. Thirteen plates were admirably engraved by another amateur, Pierre Lyonet, who was in all respects a fit companion for Trembley. It was proved that Hydra preyed upon living animals, especially upon the A nail was thus driven in a sure place. The conception of an animal was enlarged, for it was shown that an animal may branch and multiply in a way hitherto supposed to be peculiar to plants. The old connecting links between animals and plants (zoophytes, sponges, etc.) had never been really investigated; no one knew what sort of organisms formed or inhabited their plant-like skeletons. But Hydra, thanks to Trembley's description, furnished a clear example of an animal which possessed some of the attributes of a plant. Forms more ambiguous than Hydra, such as Volvox and EuglÆna, were ultimately to make the distinction between animal and plant very uncertain and shadowy. It was Hydra that gave the first clue to the structure of the zoophytes, and dispelled the false notion that corals are plants, bearing flowers, fruits, and seeds. Baer At the time when Trembley was demonstrating the asexual propagation of Hydra, Bonnet (supra, p. 45) was demonstrating the asexual propagation of aphids. Both naturalists were natives of Geneva, and both, as well as their associate Lyonet, were in a sense pupils of RÉaumur, who not only set them an admirable example, but directed their attention to promising researches and discussed with them the conclusions which might be drawn. RÉaumur's experience had seemed to confirm Leeuwenhoek's statement (supra, p. 34) that aphids produce young alive, even though no males are to be found among them; but unlucky accidents defeated his intention to confirm it by experiment, and when Bonnet asked him to suggest a piece of work RÉaumur gave him the aphid problem. Bonnet filled a flower-pot with moist earth, introduced a food-plant together with a single new-born aphid, and covered all up with a bell-jar. In twelve days the aphid produced its first young one; in a month ninety-five had been born from the same unfertilised parent. As many as five generations were obtained without the intervention of a male, each successive parent having been isolated from the moment of its birth. It was, however, discovered, apparently by Lyonet, that though viviparous reproduction without males went on regularly so long as food was plentiful, males appeared towards the end of summer, and fertilised the eggs which were destined to outlast the winter. The aphids added a new and peculiar example to the known cases of asexual propagation (plants and Hydra). Much discussion followed, but the physiology of that age (and the same is true of the physiology of our own age) was unable to reveal the full significance of the observed facts. Insects have since furnished many instances of unfertilised eggs which yield offspring. One such instance was already recorded, though neither Leeuwenhoek, RÉaumur, nor Bonnet knew of it. In the year 1701 Albrecht of Hildesheim placed a pupa in a glass vessel and forgot it. A moth hatched out and laid eggs, from which a number of caterpillars issued. Lyonet, whom we have more than once had occasion to mention, afterwards became celebrated as the author of one of the most laborious and beautiful of insect-monographs. The structure of the larva of the goat-moth was depicted by him in eighteen quarto plates, crowded with detail. The Historical or Comparative Method: Montesquieu and Buffon.About the middle of the eighteenth century we remark the introduction of a new, or almost new, method of investigation, which was destined to achieve great results. Hitherto many men had been sanguine enough to believe that they could think out or decide by argument hard questions respecting the origin of what they saw about them. It was easier, but not really more Most writers call this method historical, because history is the chief means by which it seeks to trace causes. Others call it genetic, because it goes back, whenever it can, to origins. It might also be called comparative, because it compares, not only things which are widely separated in time, but also things which are separated in space, things which differ in form or tendency because they have a common origin, and things which differ in origin because they have a common form or tendency. Whether the institutions, arts, and usages of mankind, or the species of plants and animals, are in question, the study of history, together with the comparative study of what now exists, results in increased attention to development, and this again brings to light the continuity of all natural agents and processes—continuity in time and continuity among co-existences. Since the new method has succeeded in tracing the causes of many phenomena which once seemed to obey no law, it has done much to strengthen the belief in universal causation. Down to the middle of the eighteenth century the book of Genesis had been almost unanimously accepted in Europe as the only source of information concerning the origin of the world, of man, of languages, of arts and sciences. The whole duration of the world was restricted to so brief a space that slow development Montesquieu (Esprit des Lois, 1748) was the first to exhibit on an impressive scale the power of the historical method. Natural development, determined by unalterable conditions, was with him the key to the right understanding of the past. It is well known that here and there a great thinker had before Montesquieu framed something like the same conception. The Politics of Aristotle Buffon's place in the history of science is that of one who accomplished great things in spite of weaknesses peculiarly alien to the scientific spirit. It was mainly he who, by strenuous exertions and largely at his own cost, transformed the gardens from which the king's physicians used to procure their drugs into what we now know as the Jardin des Plantes. By the untiring labours of fifty years he produced a Natural History in thirty-six volumes crowded with plates. Having won for himself a place side by side with Montesquieu and Gibbon, he employed it to direct attention to the larger questions of biology and geology. He was a pronounced freethinker, who promulgated bold views with a dexterity which saved him from condemnation by the theological tribunals. When his opinions were declared to be contrary to the teaching of the Church, he printed a conciliatory explanation, but never cancelled the passages objected to, which continued to appear in a succession of editions. His deficiencies, we must admit, were serious. He was a poor observer (partly because of short sight), and had no memory for small details. His enemies were able to taunt him with absurd mistakes, such as that cows shed their horns. He alienated the two foremost naturalists of the eighteenth century, LinnÆus and RÉaumur, by ignorant and scornful criticisms. His strong propensity to speculation, insufficiently checked by care to verify, might have brought him under the sarcastic remark of Fontenelle, that ignorance is less apparent when it fails to explain what is, than when it undertakes to explain what is not. Buffon's fame is not seriously impaired by the fact that his great work is no longer read except by those who study the course of scientific thought. Few productions of the human intellect retain their value after a hundred years, and scientific treatises become obsolete sooner than others. It is consoling to recollect that, if their energy is quickly dissipated, it is at least converted into light. In a history of biology Buffon is naturally a more important figure than Montesquieu. Buffon had imbibed evolutionary views from the ProtogÆa of Leibnitz, which in turn made use of certain hypotheses of Against all his shortcomings we must set the fact that Buffon strove to interpret the present by the past, the past by the present, geology by astronomy, geographical distribution by the physical history of the continents. One of his maxims expresses the fundamental thought of Lyell's Principles of Geology: "Pour juger de ce qui est arrivÉ, et mÊme de ce qui arrivera, nous n'avons qu'À examiner ce qui arrive." Hard-and-fast distinctions are the marks of imperfect theory. Early philosophers distinguished hot and cold, wet and dry, light and dark, male and female, as things different in kind. In later times organic and inorganic, animal and vegetable, the activities of matter and the activities of mind, have been sharply separated. But as knowledge increases these distinctions melt away; it is perceived that the extreme cases are either now connected by insensible gradations, or else spring historically from a common root. Hutton, Lyell, and their successors have made it clear that the history of the earth calls for no agents and no assumptions beyond those that are involved in changes now going on; the present is heir by unbroken descent to the past. Continuity has been established between all forms of energy. Even the chemical elements, once the emblems of independence, give indications that they too had a common origin. The nebular hypothesis, which has been steadily rendered more probable by the scientific discoveries of two centuries, traces all that can be perceived by the senses to a homogeneous vapour, and lays the burden of proof on those who believe that continuity has its limits. Every history, whether of planetary systems, or of the earth's crust, or of human civilisations, religions, and arts, is recognised as a continuous development with progressive differentiation. Amateur Students of Living Animals.A history of biology would be incomplete which took no notice of every-day observations of the commonest forms of life. Some of the best are due to the curiosity of men with whom natural history was no more than an occasional recreation. William Turner (a preacher, who became Dean of Wells), Charles Butler (a schoolmaster), Caius and Lister (physicians), Claude Perrault (a physician and architect), MÉry and Poupart (surgeons), Frisch (a schoolmaster and philologue), Lyonet (an interpreter and confidential secretary), Roesel (a miniature painter), Henry Baker (a bookseller, who gained a competence by instructing deaf mutes), White used his influence to encourage what may be called live natural history, which, as he understood it, "abounds in anecdote The many amateurs of the eighteenth century naturally demanded books written to suit them, and illustrated books with coloured plates, coming out in parts, found a ready sale. Some were devoted to insects, others to microscopic objects. In accordance with prevalent belief, the writers made a point of tracing the hand of Providence in the minutest organisms; many popular treatises were altogether devoted to natural theology. Some few of these natural history miscellanies contained original work, which has not yet lost its interest. The best is Roesel's Insecten-belustigungen (four vols. 4to., 1746-61), memorable among other things for containing the original description of Amoeba. For English readers Henry Baker wrote The Microscope Made Easy (1743) and Employment for the Microscope (1753). Intelligence and Instinct in the Lower Animals.The period with which we are now concerned (1741-1789) initiated the profitable discussion of the mental powers of animals. We are unable for lack of space to follow the investigation from period to period, and must condense into one short section whatever its history suggests. In the year 1660 Aristotelians were still discoursing about the vegetative and sensitive souls which bridged the gulf between inanimate matter and the thinking man. Descartes had tried to prove that the bodies of men and animals are machines actuated by springs like watches. Man, however, according to Descartes, possesses a soul wholly different in its properties from his body, and apparently incapable of being acted upon Swammerdam and RÉaumur, besides many naturalists of less eminence, recorded a host of observations on the activities of insects. They contributed little to the discussion except new facts, for habit led them to ascribe without reflection every contrivance to the hand of Providence or else to Nature. Some of their facts, however, made a deep impression, none more than the exact agreement of the cells of the honeycomb with the form which calculation showed to be most advantageous. Buffon The elder Huber seems to have denied to bees every trace of intelligence, but his son Pierre found it hard to go so far. In spite of examples like this, the favourite expression "blind instinct" helped to strengthen the conviction that the mental processes of animals are unsearchable. It is impossible to deny that the epithet blind is appropriate in many cases. A bird will sit an addled egg all summer, or vainly but repeatedly attempt to make its tunnel in the insufficient breadth of a mud wall (Geositta). Of course such instances do not show that all the acts of the lower animals are devoid of intelligence. Hume in 1739 and again in 1748 appealed to everyday observation of dogs, birds, and other animals of high grade. The facts seemed to him to show that animals as well as men are endowed with reason and able to draw inferences; he did not, however, credit them with the power of framing general statements, holding that experience operates on them, as on children and the generality of mankind, by "custom" alone. It is notorious that the dog and other higher animals learn by experience; Hume tells, for instance, how an old greyhound will leave the more fatiguing part of the chase to younger dogs, and place himself so as to meet the hare in her doubles. On the other hand (though Hume does not say so) man himself possesses non-educable instincts. In short, Hume sees no ground for drawing a line between the mental powers of man and those of the higher animals, though he attributes to The detailed proofs which Hume was not enough of a naturalist to furnish were at length stated with admirable clearness and force by Leroy, whose Letters on Animals form the most important contribution made to the discussion during our period. Georges Leroy (1723-1789) was lieutenant des chasses under the last French kings, and had charge of the parks at Versailles and Marly. He wrote therefore with knowledge about the wolf, fox, deer, rabbit, and dog. His pages are enlivened by many touches of nature, interesting to readers who perhaps care little about psychology. Leroy attributes to the wolf observation, comparison, judgment. The wolf must mark the height of the fold which encloses a flock, and judge whether he can clear it with a sheep in his mouth. Wolf and she-wolf co-operate artfully in the running-down of prey. Sometimes the she-wolf will draw off the sheep-dog in pursuit, thus putting the flock at the mercy of her mate. Or one of the two will chase the quarry till it is out of breath, when the other can take up the running on advantageous terms. An old fox shows knowledge of the properties of traps, and will rather make a new outlet or suffer long famine than encounter them. But when he finds a rabbit already caught, he realises that the trap has lost its power to hurt. Sheep-dogs can be educated to mind things which do not interest wild dogs, or dogs of other breeds; when, for instance, the flock is driven past a patch of wheat, the dog in charge will take care that the sheep do not damage the crop. A trained sporting-dog learns at length to trust his own judgment, even in opposition to that of his master, and sportsmen know that they must direct young dogs, but leave old ones to act for themselves. From the middle of the eighteenth century to the present day naturalists and psychologists have been labouring to distinguish instinct from intelligence. It is not hard to define well-marked examples of each, and to show that a typical instinct is congenital (not the result of a process of education or self-education), adaptive (conducive to the welfare of the organism), co-ordinated by nerve-centres (thus excluding the superficially similar behaviour of the lowest animals and all plants), actuating the whole organism (thus excluding most, if not all, reflex acts in the higher animals, as well as the wonderful adjustments effected by bone-corpuscles and other parts of organisms), and common to all the members of a species or other group (thus excluding individual aptitudes). Many naturalists entertain some form of what may be called the use-and-disuse or inherited-memory theory, supposing that the aptitudes of the offspring are influenced by the activities of the parent. Some cling to the belief that habits can be fixed and transmitted, and we must admit that the fixation and transmission of habits might explain a great deal. But all the evidence goes to prove that habits are not inherited at all, and that we must look elsewhere for the origin of instincts. Let naturalists who think differently try to account for the instincts of working bees or ants, which receive their psychical not less than their physical endowment from a long succession of ancestors, none of which worked for their living. Or let them try to explain the instances of spiders, insects, etc., which after egg-laying practise instinctive arts for the defence of their brood, standing over the eggs, carrying them about, blocking the entrance of the burrow, etc. May we not say that it is impossible for the acts of a parent to influence the congenital instincts of offspring which have already lost connection with the mother? But surely a theory of instinct breaks down which fails to account for the expedients by which the worker-bee, the worker-ant, and the spider provide for the safety of the unhatched brood or for the welfare of the community. Darwin's Origin of Species threw a new light upon The comparative psychology of small invertebrates, such as insects, is impeded by our imperfect knowledge of their nervous physiology. Introspection is here impossible; experimental physiology and pathology, which have done so much for the psychology of the higher vertebrates, almost impossible; analogy is a treacherous guide where the structures involved differ conspicuously. We have little to guide us in the psychology of insects except their behaviour, and that is often capable of a variety of interpretations. The only course is to adopt Pasteur's watchword, "Travaillons!"—the difficulties will diminish with time and labour. The Food of Green Plants.Common observation taught men in very early times that green plants draw nourishment from the soil, and Joseph Priestley, who had been proving that air is necessary both to combustion and respiration, made an experiment in 1771 to discover whether plants affected air in the same way that animals do. He put a sprig of mint into a vessel filled with air in which a candle had Priestley did not remark that the glass vessels employed in his experiments had been set in a window, and inattention to this point caused some of his attempts to repeat the experiment to fail. He was further perplexed by using vessels which had become coated with a film of "green matter," probably EuglÆna. Such vessels restored vitiated air, though no leaves were present, and when placed in the sun, gave off considerable quantities of a gas, Priestley's "dephlogisticated air" (oxygen). Hardly any oxygen was given off when the green matter was screened by brown paper. Water impregnated with carbonic acid was found to favour the production of the green matter. To us, who have been taught at school something about the properties of green plant-tissues, it seems obvious that Priestley ought to have ascertained by microscopic examination whether his "green matter" was not a living plant. But he had always avoided the use of the microscope, his eyes being weak, and after some imperfect attempts in this way he made up his mind that the green matter was neither animal nor vegetable, but a thing sui generis. Neglecting his most instructive experiments, and not waiting till he could devise new ones, or even disentangle his thoughts, he sent to the press a confused explanation, which seemed to teach that vitiated air may be restored by sunlight alone. A Dutch physician, named John Ingenhousz, who was then living in England, read Priestley's narrative and began to investigate on his own account. Without detailing his numerous experiments, we may give his own clear summary (condensed). "I observed," Ingenhousz says, "that plants have a faculty to correct bad air in a few hours; that this wonderful operation is due to the light of the sun; that it is more or less brisk according to the brightness of the light; that only the green parts of the plant can effect the change; that leaves pour out the greatest quantity of oxygen from their under surfaces; that the sun by itself has no power to change the composition of air." It will be seen that Priestley started the inquiry, devised and executed the most necessary experiments, and got excellent results. Then he lost his way, and bewildered by conflicting observations, which he was too impatient to reconcile, published a barren and misleading conclusion. Nothing was left for him but to acknowledge that Ingenhousz had cleared up all his perplexities. Nicholas Theodore de Saussure, son of the Alpine explorer, showed in 1804 that when carbon is separated from the carbonic acid of the air by green plants, the elements of water are also assimilated, a result which owes its importance to the fact that starch is a combination of carbon with the elements of water. Saussure also proved that salts derived from the soil are essential ingredients of plant-food, and that green plants are unable to fix the free nitrogen of the air; all the nitrogen which they require is obtained from the ground. We are unable to follow the history further. Though the main facts were established as early as the beginning of the nineteenth century, experimental results of The Metamorphoses of Plants.Speculations concerning the nature of the flower roused at one time an interest far beyond that felt in most botanical questions. The literary eminence of Goethe, who took a leading part in the discussion, heightened the excitement, and to this day often prompts the inquiry: What does modern science think of the Metamorphoses of Plants? Let us first briefly notice some anticipations of Goethe's famous essay. In the last years of the sixteenth century Cesalpini, taking a hint from Aristotle, tried to establish a relation between certain parts of the flower and the component layers of the stem. LinnÆus worked out the same notion more elaborately, and with a confidence which sought little aid from evidence. His wonderful theory of Prolepsis (Anticipation) need not be described, far less discussed, here. He also borrowed and adapted an analogy which had been thrown out by Swammerdam. The bark of a tree, which according to the theory of Prolepsis gives rise to the calyx of the flower, he compared to the skin of a caterpillar, the expansion of the calyx to the casting of the skin, and the act of flowering to the metamorphosis by which the caterpillar is converted into a moth or butterfly. More rational than the speculations just cited, and more suggestive to the morphologists of the future, are his words: "Principium florum et foliorum idem est" (Flower and leaf have a common origin)—which was not, however, a very novel remark in the eighteenth century. Long before LinnÆus early botanists had remarked the resemblance of sepals, petals, and seed-leaves At the very time when LinnÆus was occupied with his fanciful analogies, a young student of medicine named Caspar Friedrich Wolff, who was destined to become a biologist of great note, published a thesis which he called Theoria Generationis (Halle, 1759). This thesis marks an epoch in the history of animal embryology, but what concerns us here is that Wolff examined the growing shoot, and there studied the development of leaf and flower. He found that in early stages foliage-leaves and floral-leaves may be much alike, and thought that he could trace both to a soft or even fluid substance, which is afterwards converted into a mass of cells. It seemed to him possible to resolve the flowering shoot into stem and leaves only. Wolff's thesis, or at least that part of it which dealt with the plant, was little read and soon forgotten; his studies of the development of animals were carried further and became famous. Goethe in 1790 revived Wolff's theory of the flower, without suspicion that he had been anticipated. It is only our ignorance, he said, when the fact came to his knowledge, that ever deludes us into believing that we have put forth an original view. As soon as he realised the true state of the case, he spared no pains to do Wolff full justice. The aim of Goethe's Metamorphoses of Plants was to determine the Idea or theoretical conception of the plant, and also to trace the modifications which the Idea undergoes in nature. These two inquiries constituted what he called the Morphology of the plant, a useful, nay, indispensable term, which is still in daily use. He thought that he could discover in the endless variety of LinnÆus's exposition of the nature of the flower had been read attentively by Goethe, who must have remarked that the conversion of organs to new uses was there described as a metamorphosis. That word had been, long before the time of LinnÆus, appropriated to a particular kind of change—viz., an apparently sudden change occurring in the life-history of one and the same animal. It was therefore unlucky that Goethe should have been led by the example of LinnÆus to employ the word in the general sense of adaptation to new purposes. He did not, however, expressly compare flower-production with the transformation of an insect, as LinnÆus had done. The reception of Goethe's Metamorphosen der Pflanzen Then came the discoveries of Hofmeister, followed by Darwin's Origin of Species. Naturalists soon ceased to put the old questions, and the old answers did not satisfy them. Wolff and Goethe had generalised the flowering plant until it became a series of leaf-bearing nodes alternating with internodes, but no such abstract conception could throw light upon the common ancestor of all the flowering plants, nor upon the stages by which the flowering plant has been evolved, and it was these which were now sought. Hofmeister brought to light a fundamental identity of structure in the reproductive organs of the flowering plants and the higher cryptogams. There has since been no doubt in what group of plants we must seek the ancestor of the flowering plant. It must have been a cryptogam, not far removed from the ferns, and furnished with sporophylls—i.e., leaf-like scales, on which probably two kinds of sporangia, lodging male and female spores respectively, were borne. The careful investigation of the fossil plants of the coal measures has brought us still nearer to the actual progenitor. Oliver and Scott The discovery of a fossil plant which makes so near an approach to the cryptogamic ancestor of all the flowering plants may remind us how little likely it was that the ideal plant of Wolff and Goethe, consisting of leaves, stem, and other vegetative organs, but without true reproductive organs, should fully represent the type from which the flowering plants sprang. No plant so complex as a fern could maintain itself indefinitely without provision for the fertilisation of the ovum; the only known asexual plants are of low grade, and, it may be, insufficiently understood. What substratum of plain truth underlies the doctrine of the metamorphoses of plants? Botanists would agree that all sporophylls, however modified, are homologous or answerable parts. Carpels and stamens are no doubt modified sporophylls. Petals are sometimes, perhaps always, modified stamens, and therefore modified sporophylls also. We must not call a sporophyll a leaf, for it contains a sporangium of independent origin, and the sporangium is the more essential of the two. The common origin of foliage-leaf, bract, perianth-leaf, sporophyll (apart from the sporangium), and seed-leaf is unshaken. We may picture to ourselves a plant clothed with nearly similar leaves, some of which either bear sporangia or else lodge sporangia in their axils. Part of such a primitive flowering plant might retain its vegetative function and become a leafy shoot, while another part, bearing crowded sporophylls, might yield male, female, or mixed cones. From an ancestor thus organised any flowering plant might be derived. But the chief wonder of the theory of Metamorphoses—viz., the derivation of stamen and pistil from mere foliage-leaves—disappears. Anther and ovule take their real origin from the sporangium, whose supporting leaf is only an accessory. The chief steps by which the morphology of the flowering plant has been attained are these:—Cesalpini (1583), followed by several other early botanists, recognized the fundamental identity of foliage-leaf, perianth-leaf, and seed-leaf. LinnÆus (1759) added stamen and carpel to the list, identifications of greater interest, but only partially defensible. Wolff (1759) justified by similarity of development the recognition of floral organs as leaves. Goethe (1790) traced structural similarity, transitions, and retrogression in leaves of diverse function. Hofmeister (1849-57) showed a relationship between the flowering plant and the higher cryptogams. Oliver and Scott (1904), inheriting the results of Williamson's work, discovered a carboniferous seed-bearing plant, one of a large group intermediate between ferns and cycads. It is now possible to explain the resemblance of the various leaf-like appendages of the flowering plant by derivation either from the leaves or the sporophylls (the latter not being wholly leaves) of some extinct cryptogam, which was either a fern or a near ally of the ferns. Early Notions about the Lower Plants.The fathers of botany neglected everything else in order to concentrate their attention upon the flowering plants, from which very nearly all useful vegetable products were derived. The lack of adequate microscopes rendered it almost impossible to investigate the structure and life-history of ferns, mosses, fungi, and algÆ until the nineteenth century. As late as the time of LinnÆus it was possible to maintain that they developed spontaneously, though the great naturalist himself called them Cryptogamia, thereby expressing his conviction that they reproduce their kind like other plants, but in Ferns.—Even before the age of LinnÆus it was known that little ferns spring up around the old ones, and that a fine dust can be shaken from the brown patches on the back of ripe fern-leaves. The dust was reputed to be the seed of the fern, and in an age which believed in magic the invisibility of fern-seed made it easy to suppose that the possessor of fern-seed would become invisible also. When the microscope began to be applied to minute natural objects, the brown patches of the fern-leaf were closely examined. William Cole of Bristol (1669), Malpighi, Grew, Swammerdam, Leeuwenhoek, and others, found the stalked capsules (sporangia), their elastic ring and the minute bodies (spores) lodged within them; it seemed obvious to call the capsules ovaries and the spores seeds. Some time in the latter part of the seventeenth century Robert Morison, professor of botany at Oxford, who died in 1683, sowed spores of the harts-tongue fern, and next year got an abundant crop of prothalli, which he took to be the cotyledons. A little later, when it had been proved that flowering plants possess male and female organs, diligent search was made for the stamens and pistils of ferns and mosses, which of course could not be found, though identifications, sometimes based upon a real analogy, were continually announced. Late in the eighteenth century one John Lindsay, a surgeon in Jamaica, who was blest with leisure and a good microscope, repeated the experiment of Morison, which seems to have been almost forgotten. Having remarked that after the rains young ferns sprang up in shady places where the earth had been disturbed. Mosses.—LinnÆus observed that the large moorland hair-moss (Polytrichum) is of two forms, only one of which bears capsules, and further that in dry weather the capsules emit masses of fine dust. No further progress was made until 1782, when Hedwig, in a memoir of real merit, described the antheridium and archegonium of the moss, and traced the capsule to the archegonium. Interpreting the organs of the moss by those of the flowering plant, he called the antheridia anthers, the capsule was a seed-vessel, the spores were seeds, and the green filament emitted by the germinating spore a cotyledon. Such misinterpretations were then inevitable. Fungi.—Micheli in 1729 found the spores of several fungi, germinated them, and figured the product. The figures show the much-branched filament (mycelium) which burrows in the soil and constitutes the vegetative part of the fungus, and also here and there a pileus (mushroom, toadstool, &c.), which is the fructification springing out of the mycelium. His account comprises the best part of what is known down to the present time of the reproduction of that group of fungi to which the mushroom belongs. AlgÆ.—Some early observers (RÉaumur among the rest) studied the enlarged and fleshy branches of brown seaweeds, and discovered the seed-like spores. This scanty knowledge of the life-history of cryptogams sufficed until the nineteenth century, when the study was resumed with better microscopes and in a far more connected way, with results of the highest interest and importance (see below, p. 108). |