Chapter III: Part 3

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 man a power of demonstrative reasoning to which animals do not attain. In this he substantially agrees with Aristotle,[28] who maintained that in animals the germs of the psychical qualities of the man are evident, though, as in the child, they are undeveloped. Hume's teaching also accords with modern views; comparative anatomy, for instance, "is easily able to show that, physically, man is but the last term of a long series of forms, which lead by slow gradations from the highest mammal to the almost formless speck of living protoplasm, which lies on the shadowy boundary between animal and vegetable life."[29]

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).[30] In the same way it is easy to point out clear differences between a bird and a tree. But just as a definition which shall separate every animal from every plant has hitherto been sought in vain, so it has hitherto been impossible to frame a definition which while including all instincts shall admit no case of reflex action or intelligence. The most ambiguous cases of all are perhaps to be found in insects, where, as will shortly be explained, our information is ill-fitted to support precise distinctions.

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 instinct by showing that natural selection can operate on the subtlest modifications. It can discriminate shades of hardiness to climate, shades of intellectual acuteness, or shades of courage. It can intensify qualities which appear only in adults past bearing or in individuals congenitally incapable of propagation. Human selection, though a blunt tool in comparison with natural selection, can originate a bold and hardy race of dogs, or showy double flowers incapable of producing seed. In the second case fertile single flowers continue the race, as in the garden Stock. Darwin pointed out that the barren double flowers of the Stock answer to the workers of social bees and ants, the fertile single flowers to the functional males and females. Every modification that works to the advantage or disadvantage of the race, whether we classify it as physical, intellectual, or moral, gives scope for the operation of natural selection.

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 that sunlight is necessary to their health. In the age of Galileo a Belgian physician and chemist, Van Helmont, endeavoured to pursue the subject by experiment. He planted the stem of a live willow in furnace-dried earth, which was enclosed in an earthen vessel. Rain-water or distilled water was supplied when necessary, and dust excluded by a perforated lid. The loss of weight due to the falling-off of leaves was neglected. In the course of five years the tree was found to have increased to more than thirty times its original weight; Van Helmont concluded that this increase was due to water only. Malpighi (1671), being guided mainly by his microscopic studies of the anatomy of the stem and leaf, taught that moisture absorbed by the roots ascends by the wood, becoming (apparently at the same time) aerated by the large, air-conducting vessels; that it enters the leaves, and is there elaborated by evaporation, the action of the sun's rays, and a process of fermentation; lastly, that the elaborated sap passes from the leaves in all directions towards the growing parts. It will be seen that this explanation, though incomplete, makes a fair approximation to the beliefs now held; for more than a hundred years after Malpighi's day less instructed opinions were commonly held. Hales (1727) recognised that green plants are largely nourished at the expense of the atmosphere; he dwelt also on the action of the leaves in drawing water from the soil, and in discharging superfluous moisture by evaporation.

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 burned out, and after ten days found that a candle would now burn perfectly well in the same air. Air kept without a plant, in a glass vessel immersed in water, did not regain its power of supporting combustion. Balm, groundsel, and spinach were found to answer just as well as mint. Air vitiated by the respiration of mice was restored by green plants as readily as air which had been vitiated by combustion.

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 scientific and practical interest have never ceased to accumulate down to the present time.

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 to foliage-leaves; Cesalpini has a common name for all (folium).

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 the organs of the flowering plant one structure repeated again and again, which gradually attained, as by the steps of a ladder, what he called the crowning purpose of nature—viz., the sexual propagation of the race. This fundamental structure was the leaf. The proposition that all the parts of the flower are modifications of the leaf he defended by three main arguments—viz., (1) the structural similarity of seed-leaves, foliage-leaves, bracts, and floral organs; (2) the existence of transitions between leaves of different kinds; and (3) the occasional _retrogression_, as he called it, of specially modified parts to a more primitive condition. These lines of argument were illustrated by many well-chosen examples, the result of long and patient observation. Goethe did not, however, fortify his position by the likeness of developing floral organs to developing foliage-leaves, which had been Wolff's starting-point. He arrived independently at Wolff's opinion that the conversion of foliage-leaves into floral organs is due to diminished nutrition.

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_ was at first cold, but the doctrine which it enforced gradually won the attention of botanists, and by 1830 he was able to show that it had been accepted by many good judges.

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[31] have pointed out that the carboniferous Lyginodendron, though showing unmistakable affinity with the ferns, bore true seeds, as a pine or a cycad does. Many other plants of the coal measures are known to have combined characteristics of ferns with those of cycads, while some of them, like Lyginodendron, crossed the frontier, and became, though not yet flowering plants, at least seed-bearers.

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 a way so far not understood. Gaertner, a contemporary of Linnæus, pointed out one important respect in which the spores of cryptogams differ from the seeds of flowering plants, viz. that they contain no embryo.

_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. it occurred to him to mix the fine brown dust from the back of a fern-leaf with mould, sow the mixture in a flower-pot, and watch daily to see what might come up. About the twelfth day small green protrusions were observed, which enlarged, sent down roots, and formed bilobed scales, out of which young ferns ultimately grew. In 1789 Sir Joseph Banks, who was reputed to be the best English botanist of the day, asked Lindsay's help in sending West Indian ferns to Europe. Lindsay replied that it would be easier to send the seed, and that the seed would grow if properly planted. This was new to Banks, who demanded further information. Lindsay then prepared a short illustrated paper, which Banks communicated to the newly formed Linnean Society. It will be seen that Lindsay was able to add nothing of much importance to what Morison had ascertained a century before. The spores were still identified with seeds, the prothallus was still a cotyledon, and for years to come botanists continued to seek anthers on fern-leaves. At this point we suspend for a time the history of the discovery (see below, p. 108).

_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).

[15] Reden, Vol. I., pp. 109, 154.

[16] _Traité d'Insectologie_, première partie. Two vols.12 mo. Paris, 1745.

[17] In circles untouched by general European thought such beliefs lasted much later. Sir Francis Galton (_Memories of My Life_, p. 67) says: "The horizon of the antiquarians was so narrow at about the date (1840) of my Cambridge days that the whole history of the early world was literally believed, by many of the best-informed men, to be contained in the Pentateuch. It was also practically supposed that nothing more of importance could be learnt of the origin of civilisation during classical times than was to be found definitely stated in classical authors."

[18] "If anything could disentitle Montesquieu's _Esprit des Lois_ to the proud motto, _Prolem sine matre creatam_, it would be its close relationship to the Politics." (A. W. Benn's _Greek Philosophers_, Vol. II., p. 429.)

[19] For an account of other early hypotheses of the same kind the reader may refer to Edward Clodd's _Pioneers of Evolution_.

[20] _Life and Letters_, Vol. II., p. 212.

[21] _Origin of Species_, ed. i., p. 484.

[22] White uses _anecdote_ in the old sense, meaning by it a piece of unpublished information.

[23] Réaumur, _Hist. des Insectes_, Vol. V., Mém. viii.

[24] Darwin, _Origin of Species_, chap. vii.

[25] Vol. III., Mém. iv.

[26] _Hist. Nat._, Vol. IV.

[27] The first edition of the _Nouvelles Observations sur les Abeilles_ (1792) was the work of François Huber alone; the second (1814) was prepared by Pierre with the co-operation of his father, and is here credited to the son.

[28] _Hist. Animalium_, VIII., i.

[29] Huxley's _Hume_, chap. v. Some few naturalists, who are entitled to respectful attention, such as Father Wasmann, author of _The Psychology of Ants_, do not even now receive the conclusions of Hume.

[30] Lloyd Morgan, _Habit and Instinct_, Introduction.

[31] _Phil. Trans._, 1904.

PERIOD IV.

1790-1858

Characteristics of the Period.

The first French republic and the first French empire were associated with a great outburst of scientific energy. French mathematics, astronomy, and physics were pre-eminent. England suffered from isolation during the continental war, but Davy, Young, the Herschels, Watt (now past his prime), Dalton, and William Smith supported the scientific reputation of their country. In Germany this was the age of Goethe and Schiller; Alexander von Humboldt was prominent among the scientific men of Prussia. The forty years' peace, during which reaction prevailed in many parts of Europe, was in England and America a time of steady growth and progress.

Sprengel and the Fertilisation of Flowers.

Conrad Sprengel, an unsuccessful schoolmaster who lived in a Berlin attic and got his bread by teaching languages or whatever else his pupils wished to learn, wrote a book which marks an epoch in the study of adaptations. This was his _Secret of Nature Discovered_, which appeared in 1793. Half a century passed before its merit was recognised by any influential naturalist; even then the recognition was private, and never reached the author, who had died long before. There was no striking of medals, no jubilee-celebration, nothing more than this, that Robert Brown recommended the book to Charles Darwin, who found in it, as he says, "an immense body of truth."

In 1787 Sprengel had remarked that the bases of the petals of Geranium silvaticum are beset with long hairs. Persuaded that no natural structure can be devoid of meaning, Sprengel asked what purpose these hairs might serve. A honey-gland in their midst suggested that they might protect the honey by keeping off the rain, which easily enters this shallow flower. Other honey-secreting flowers were found to possess mechanisms adapted to the same end. His first question suggested a second: Why should flowers secrete honey?

Malpighi had described the honey-glands of crown-imperial (1672), and had seen that the honey must be secreted by the petals, and not deposited from the atmosphere, according to the notion then current. Kölreuter (1761) had showed that insects may effect the pollination of flowers. Linnæus (1762) had given the name of _nectary_ to the honey-gland. He thought that the honey served to moisten the ovary, though he knew of staminate flowers furnished with nectaries. He also threw out the alternative conjecture that the honey is food for insects, which disperse the pollen by their wings. Sprengel improved upon all his predecessors, and made it clear that transference of pollen is the main purpose of the honey in flowers. He was put on the right track by the study of a forget-me-not flower. Here he found the honey protected from rain by the narrowness of the corolla-tube, whose entrance was almost closed by internal protuberances. The protuberances were distinguished by their yellow colour from the sky-blue corolla, and this conspicuous colouration led Sprengel to infer that insects might be thereby induced to seek for the store of honey within. He tested his conjecture by examining other honey-bearing flowers, and soon collected many instances of spots, lines, folds, and ridges, which might not only make insects aware of hidden stores of honey, but guide them to the exact place. Contrivances of the most diverse kinds, but all tending to invite the visits of insects and utilise them for the benefit of the plant, rewarded Sprengel's continued inquiries. He found that night-flowering plants, which could derive no advantage from coloured patterns, often have large white corollas, easily discerned in a faint light, and that these flowers give out an odour attractive to nocturnal insects. He found that the pollen-masses of an orchis are actually removed by large insects, though here no honey could be detected in the flower. Sprengel's fertility in probable conjecture is shown by his explanation of this puzzling case; he suggested that the orchis is a sham honey-bearer (Scheinsaftblume), which attracts insects by assuming the conspicuous size and coloration found in most honied flowers. Darwin suspected, and Herman Müller proved, that though the spur of the orchis-flower is empty, it yields when pierced a fluid attractive to bees and other insects. Sprengel discovered too how insects get imprisoned in the corolla of an Aristolochia, whose reflexed hairs allow small flies to creep in, but effectually prevent their escape until they have fertilised the pistils, when the hairs relax. These are only specimens of a multitude of adaptations which fill the book.

Sprengel insists upon the study of flowers under natural conditions; he could never have made out by the examination of plucked flowers how Nigella is fertilised. Flies with attached pollen-masses, which he found in spiders' nests, gave him the hint as to the way in which the fertilisation of orchids is effected. Definite questions must be put if observation is to be profitable. What is the use of honey to the plant—of this coloured spot—of these hairs? He notes the peculiarities of wind-fertilised and insect-fertilised flowers, the relative abundance of the pollen, the form of the stigma, the presence or absence of honey, the size, colour, and scent of the corolla. Here is a pretty illustration from his pages. Pluck a branch of hazel, aspen, or alder, with unexpanded catkins, and also one from the male sallow; place them in water, and keep them in a sunny window until the anthers are ripe. A vigorous puff will then discharge a cloud of pollen from the _wind-fertilized_ catkins, but none from the _insect-fertilised_ catkin of the sallow. What Linnæus said about the flowers of trees appearing before the leaves, in order that the pollen may more easily reach the stigmas, holds good, Sprengel remarks, only of wind-fertilised trees. The lime, which is insect-fertilised, flowers in the height of summer, when all the branches are crowded with leaves.

Sprengel left it to later biologists to complete his discovery. "That wonderfully accurate observer, Sprengel," says Darwin,[32] "who first showed how important a part insects play in the fertilisation of flowers, called his book _The Secret of Nature Displayed_; yet he only occasionally saw that the object for which so many curious and beautiful adaptations have been acquired, was the cross-fertilisation of distinct plants; and he knew nothing of the benefits which the offspring thus receive in growth, vigour, and fertility." Not even

Cuvier and the Rise of Palæontology.

If this historical sketch had been prepared within a few years of the death of Cuvier, it would no doubt have held him up as the greatest of zoologists and comparative anatomists. Nor would it have been hard to find reasons for such a verdict. His _Règne Animal_ extended and corrected the zoological system of Linnæus; his comparative anatomy, and especially his comparative osteology, were far ampler and more exact than anything that had been attempted before. It would not have been forgotten, moreover, that he was the practical founder of the new science of palæontology.

At a later time, say in the sixties and seventies of the nineteenth century, when the Origin of Species controversy was in full blast, any estimate of Cuvier by an evolutionist would have been much less laudatory. Cuvier had actively opposed that form of evolution which had been brought forward in his day, and with such power as to close the discussion for a time. The assailants of the Origin of Species found his refutation of unity of type and progressive development adaptable to the new situation, and the reasoning which had pulverised Geoffrey St. Hilaire was brought out again in order to pulverise Darwin. Then the supporters of Darwin found it necessary to show that Cuvier was by no means infallible. This they were able to do without introducing matter foreign to the main question, for Cuvier's exposition of fixity of species, of the principles of classification and of the process of extinction, were entirely opposed to the beliefs not only of Darwin, but of Lyell and the whole school which stood out for historical continuity, treated history of every kind as a process of development, extended almost without limit the duration of life on the earth, and enforced the obvious but neglected truth that results of any magnitude whatever may proceed from small causes operating through a sufficient length of time.

Darwin's main contentions are now accepted by the scientific world, and Cuvier's hostility to particular forms of evolution has become a mere historical episode of no lasting importance. Angry disputes concerning the weight of his authority are at an end; he is not to be blamed because thirty years after his death he was set up as judge of a cause which he had not heard. We are now ready to make fair allowance for the time in which his lot was cast—an age when geology, embryology, palæontology, and distribution were mere infants, some of them hardly yet born. We can also admit without reserve the incompetence of certain of Cuvier's antagonists, and justify the severity with which he treated unity of type as stated and defended by Geoffroy St. Hilaire. Now that the dust of controversy has settled, we are chiefly concerned to inquire: What of all Cuvier's work has proved to be really permanent? His zoology and his comparative anatomy have had to be completely re-cast, partly because of the new light thrown on them by embryology and the doctrine of descent with modification. His studies of extinct vertebrates, however, called into existence a new science, the science of Palæontology,[33] and it is mainly this which gives him a lasting and honoured place in the history of biology.

At the end of the eighteenth century it had been rather grudgingly admitted that some few animals were actually extinct. Buffon was able to quote as indubitable examples the mammoth and the mastodon. Their occurrence in countries unknown to the ancients, such as Siberia and North America, disposed of the explanation long clung to by the learned—viz., that their bones were the remains of elephants which had been led about by the Roman armies, while their large size and the ease with which they can be recognised rendered it highly improbable that they still survived anywhere on the surface of the globe.

It was therefore natural that Cuvier's first study in palæontology should relate to extinct elephants. He compared and distinguished several species, showed that they were distinct from the existing Asiatic and African species, a fact which had escaped the notice of Pallas, and argued from the well-known case of a Siberian mammoth preserved in ice and frozen mud with hardly any decomposition that it must have been overwhelmed by a sudden "revolution of the earth." Whatever we may think of Cuvier's geology, his comparisons of all known elephants, recent and fossil, introduced a new standard of exactness into these inquiries. From this beginning he went on to study all the extinct vertebrates which he could discover in public or private collections. By 1821 he had published elaborate and well-illustrated descriptions of near a hundred extinct animals, an extraordinary output for one investigator.

The most remarkable of his palæontological discoveries were made at home, in the lower tertiary rocks which underlie the city of Paris. He proved that in the valley of the Seine a large population of animals, all now extinct, had formerly flourished. None of these discoveries impressed his contemporaries more than the celebrated case of the fossil opossum. The bones were imbedded in a slab of gypsum, and were at first imperfectly exposed. The lower jaw, however, exhibited a peculiarity of marsupial or pouched animals, for its angle had an inwardly projecting shelf, not found in other quadrupeds. The opossums, like all marsupial animals, bear on the front of the pelvis two long bones, which support the pouch. These were as yet concealed, and Cuvier delayed clearing them until he had summoned friends, some of whom may have been sceptical about the possibility of reasoning with certainty from anatomical data. Warning them what to expect, he removed with a sharp tool the film of stone, and revealed the long and slender marsupial bones.[34] The ancient existence of marsupials in France was then a striking and almost incredible fact; increase of knowledge has not lessened its interest, though it has abated some of the wonder.

The fossil ungulates (hoofed quadrupeds) of the Paris basin taxed Cuvier's patience and skill to the utmost. In the tiresome work of piecing together a multitude of imperfect skeletons he set an example to all future palæontologists. That he drew general conclusions which we are unable to accept, and failed to draw conclusions which seem obvious to us, will surprise nobody whose reading has taught him how unprepared were the biologists of that age to handle great questions concerning the origin and extinction of races. Cuvier recognised among the fossils of the Paris quarries the bones of two genera of ungulates very different from any of recent times. One resembled the rhinoceros, tapir, and horse in being odd-toed; this he called Palæotherium. Another had the hind-foot even-toed, like a ruminant, though the fore-foot, with which he was imperfectly acquainted, showed points of resemblance to the other group. How cautiously he did his work may be gathered from the fact that he spent fifteen years upon the collection of facts before he attempted to restore these extinct forms, though almost every bone in their bodies had during that time passed through his hands.

The great interest of these fossil ungulates to the modern biologist is that they are relatively primitive types of the order. Palæotherium is not far from the ideal common ancestor of the rhinoceros, tapir, and horse; Anoplotherium not altogether unlike the ideal common ancestor of the hippopotamus, the swine, and the ruminants. It has been suspected that Cuvier was less obstinately devoted to the tenet of fixity of species than he was willing to admit in public. Whatever his private leanings may have been, he stood out resolutely for cogent proofs of transmutation. When it was contended that the Palæothere might have been the remote ancestor of existing ungulates, he demanded that the intermediate links should be produced. His demand could not be met till many years later, though intermediate forms between the Palæothere and the horse have since been furnished in abundance. Reserve about far-reaching deductions was surely wise at a time when plausible speculation was rife, and we ought not to judge Cuvier severely for having aspired to a rigour unattainable in a natural science, and certainly not always observed by himself. He hoped to see biology become as exact as astronomy. The hope may have been chimerical, but emphasis on this side was not altogether out of place in the generation of Geoffroy St. Hilaire and Oken.

If the great master who laid the foundations of palæontology could revisit the scene of his former labours, he would find that many strange things had happened since the appearance of his _Ossemens Fossiles_. He would perhaps be stupefied at first to discover how little is now made of the Revolutions of the Earth, the proofs of which had seemed to him unimpeachable, while the conjectures about the development of new races, which in his own day had been almost negligible, have proved to be anticipatory of fundamental biological truths. The first shock over, one can imagine the zest with which he would strive to combine the familiar facts into a body of new doctrine. The ungulates, recent and fossil, would of course interest him particularly. He would recognise the gradations of structure which run through the whole order, branching and crossing in all directions; gradation in the number of the toes, in the rearing of the body more and more upon the toe-tips, in the progressive complication of the teeth. One chain of examples would lead from the shallow, tuberculate molar of the pig to the molar of the horse or ruminant, deep and massive, with crescentic enamel-folds; another would illustrate the gradual development of tusks from ordinary incisors or canines; a third series would show the steps by which the primitive ungulate dentition became reduced to the dentition of the elephant, with only a single pair of incisors, enlarged into tusks several feet long, with no canines but molars of great weight, complicated by extreme folding. It would surprise and delight him to compare the almost insensible steps by which his own Palæothere can be seen to pass into the modern horse. Then we can imagine how our regenerate Cuvier would draw nearer and nearer to the common ancestor of the whole group, a five-toed, plantigrade ungulate, with the full dentition of forty-four unspecialised teeth, and how readily he would admit that Phenacodus, both in its structure and its geological horizon, was just the common ancestor that theory required. The proofs of intermediate stages between ancient and modern ungulates which he had once called for in vain, he would now find ready to his hand. It might well seem that the history of the ungulates, with all its modern expansions, would suffice to occupy even his unparalleled energy. He would see with delight how the palæontology which he had been the first to treat as a science has enlarged the comparative anatomy of which also he was so great a master. He would cheerfully admit that both yield proofs of that doctrine of descent with modification which a hundred years ago seemed to him so questionable.

Chamisso on the Alternation of Generations in Salpa.

Trembley (see p. 57) had shown that Hydra, though an animal, multiplies by budding like a plant. He got indications, upon which he did not altogether rely, that it also propagated by eggs, and ten years later (1754) this supposition was confirmed by Roesel, who figured the egg, though he was unable to demonstrate that a young Hydra issues from it; subsequent inquiry has placed the fact beyond doubt. In 1819 Chamisso announced that Salpa, a well-known Tunicate which abounds at the surface of the sea, exhibits a regular alternation of the two modes of increase, the egg-producing form being succeeded by a budding form, the budding form by the egg-producing form, and so on indefinitely. Sars a few years later showed that the common jelly-fish Aurelia also propagates by eggs and buds alternately. Here the familiar swimming disks, which are of two sexes, produce eggs from which locomotive larva issue. The larva at length settles down and takes a Hydra-like form. It pushes upwards an ascending column, which divides transversely and forms a pile of slices, each destined to become a free, sexual Aurelia. The alternation of generations may be regarded as resulting from the introduction of budding into the early stage of a life-history which culminates in sexual reproduction, much as if a caterpillar were to divide repeatedly and form more caterpillars, each of which ultimately became a moth. The case which has been given as an illustration actually occurs in nature. A parasitic caterpillar, that of Encyrtus, divides while still an embryo, so that one egg produces several moths.[35] Many other cases of alternation have since been found among animals, and it seems to be the rule among plants.

Alternation of generations may be complicated by association with transformation, by the omission of stages usual in the class, and by budding-out from one part instead of from the whole body. In particular cases the complication becomes so great that biological language breaks down under it. Such terms as _generation_, _individual_, _organ_, _larva_, _adult_, cannot always be used consistently without either being strained or artificially limited.

Baer and the Development of Animals.

The curiosity of the ancient Greeks led them to look for the chick within the egg, and Aristotle mentions the beating of the heart as a thing which might be observed in a third-day embryo. After the revival of science Fabricius of Acquapendente figured the chief stages of development, from the first visible rudiments to the escape from the egg-shell. Harvey, the discoverer of the circulation, not only studied the developing chick, but took advantage of the rare opportunity of dissecting breeding does from the royal parks. His treatise on Generation is unfortunately impaired by Aristotelian philosophy, and some of the theories there set forth gave much trouble to Swammerdam. The oft-cited maxim "Omne vivum ex ovo" does not occur exactly in this form in Harvey's writings,[36] nor does it fairly state his own belief. Those who read his _De Generatione_ will see that his knowledge was insufficient to justify so wide a generalisation; on this head it is enough to mention that he was persuaded of the production of insects without parents from putrefying matter.[37]

Malpighi was the first to apply the microscope to the embryonic chick. His figures are surprisingly full of interesting detail, and so far in advance of their age that they long failed to produce their due effect. On one point Malpighi unconsciously led naturalists astray for a hundred years or more. On examining a fowl's egg which he supposed to be unincubated, he discovered within it an early embryo. From this he concluded that the embryo _pre-exists_ in the egg, like a plant-embryo in a seed. He mentions one circumstance which makes everything intelligible. The egg was examined in August, during a time of great heat, and the Italian summer no doubt started development, like the hot sand of Aden, in which Chinamen hatch their eggs. Swammerdam too enforced the same belief in pre-existing germs. From the fact that the butterfly can be revealed by opening the skin of a full-fed caterpillar he inferred (quite contrary to the opinion which he expresses elsewhere) that one animal had formed inside another. This led him to say that there is no such thing as generation in nature, but merely the expansion of germs which lie enclosed one within another. By his theory he explained how Levi could pay tribute to Melchizedek before he was born, and how the sin of Adam can be laid to the charge of all his posterity. The belief in the pre-existence of germs was first shaken by Caspar Wolff (see p. 81), who examined unincubated eggs but found no germ which could be detected by the histological methods then employed.

Swammerdam's _Biblia Naturæ_ contains useful figures of early and late tadpoles; in particular, he describes a stage in which the body is entirely composed of rounded "lumps" or "granules," the _cells_ of modern biology.

Early in the nineteenth century Pander and Baer, both of whom were pupils of Döllinger, a teacher of extraordinary influence, gave a new impetus to the study of development. Pander (1817-8) published an account of the early stages of the chick, illustrated by beautiful plates by D'Alton. Baer (1828-37) carried the work much further, not only greatly extending the knowledge of the developing chick, but discovering the mammalian ovum (1827), and announcing generalisations which down to 1859 were the most luminous that embryology had ever furnished; we may call him the founder of comparative embryology. He shows that development may supply decisive indications of the zoological position of animals; it teaches, for instance, that insects are of higher grade than arachnids or crustaceans, and that amphibians ought not to be united with reptiles. He describes the development of an animal as a process of differentiation, the general becoming special, and the homogeneous heterogeneous; differentiation is, he remarks, the law under which not only animals but solar systems develop. He maintains that the embryo, though gradually attaining complexity, makes no transition to a different type—_e.g._, the vertebrate is never in any stage anything but a vertebrate. All animals, he believes, are probably at first similar, and take the form of a hollow sphere (the _gastræa_ of modern embryology). There are, he says, no new formations in nature; all is conversion. When he comes to speak of the pharyngeal clefts of mammals and birds, recently discovered by Rathke, he remarks that their correspondence with the gill-clefts of fishes is obvious. We wonder what is coming next, but our curiosity is not gratified by any memorable deduction. Neither here nor in his miscellanies (_Reden_), published nearly fifty years later, does he admit that mammals and birds can have descended from gill-breathing vertebrates. If we are inclined to hint that Baer, having gone so far, might well have gone a little farther, it is only fair to recollect that every leader in science is more or less open to the same reproach.

The Cell Theory.