Schleiden freely acknowledged his indebtedness to Brown for first knowledge of the nucleus, but he soon carried his studies of that structure far beyond those of its discoverer. He came to believe that the nucleus is really the most important portion of the cell, in that it is the original structure from which the remainder of the cell is developed. Hence he named it the cytoblast. He outlined his views in an epochal paper published in Muller's Archives in 1838, under title of "Beitrage zur Phytogenesis." This paper is in itself of value, yet the most important outgrowth of Schleiden's observations of the nucleus did not spring from his own labors, but from those of a friend to whom he mentioned his discoveries the year previous to their publication. This friend was Dr. Theodor Schwann, professor of physiology in the University of Louvain.

At the moment when these observations were communicated to him Schwann was puzzling over certain details of animal histology which he could not clearly explain. His great teacher, Johannes Muller, had called attention to the strange resemblance to vegetable cells shown by certain cells of the chorda dorsalis (the embryonic cord from which the spinal column is developed), and Schwann himself had discovered a corresponding similarity in the branchial cartilage of a tadpole. Then, too, the researches of Friedrich Henle had shown that the particles that make up the epidermis of animals are very cell-like in appearance. Indeed, the cell-like character of certain animal tissues had come to be matter of common note among students of minute anatomy. Schwann felt that this similarity could not be mere coincidence, but he had gained no clew to further insight until Schleiden called his attention to the nucleus. Then at once he reasoned that if there really is the correspondence between vegetable and animal tissues that he suspected, and if the nucleus is so important in the vegetable cell as Schleiden believed, the nucleus should also be found in the ultimate particles of animal tissues.

Schwann's researches soon showed the entire correctness of this assumption. A closer study of animal tissues under the microscope showed, particularly in the case of embryonic tissues, that "opaque spots" such as Schleiden described are really to be found there in abundance—forming, indeed, a most characteristic phase of the structure. The location of these nuclei at comparatively regular intervals suggested that they are found in definite compartments of the tissue, as Schleiden had shown to be the case with vegetables; indeed, the walls that separated such cell-like compartments one from another were in some cases visible. Particularly was this found to be the case with embryonic tissues, and the study of these soon convinced Schwann that his original surmise had been correct, and that all animal tissues are in their incipiency composed of particles not unlike the ultimate particles of vegetables in short, of what the botanists termed cells. Adopting this name, Schwann propounded what soon became famous as his cell theory, under title of Mikroskopische Untersuchungen uber die Ubereinstimmung in der Structur und dent Wachsthum der Thiere und Pflanzen. So expeditious had been his work that this book was published early in 1839, only a few months after the appearance of Schleiden's paper.

As the title suggests, the main idea that actuated Schwann was to unify vegetable and animal tissues. Accepting cell-structure as the basis of all vegetable tissues, he sought to show that the same is true of animal tissues, all the seeming diversities of fibre being but the alteration and development of what were originally simple cells. And by cell Schwann meant, as did Schleiden also, what the word ordinarily implies—a cavity walled in on all sides. He conceived that the ultimate constituents of all tissues were really such minute cavities, the most important part of which was the cell wall, with its associated nucleus. He knew, indeed, that the cell might be filled with fluid contents, but he regarded these as relatively subordinate in importance to the wall itself. This, however, did not apply to the nucleus, which was supposed to lie against the cell wall and in the beginning to generate it. Subsequently the wall might grow so rapidly as to dissociate itself from its contents, thus becoming a hollow bubble or true cell; but the nucleus, as long as it lasted, was supposed to continue in contact with the cell wall. Schleiden had even supposed the nucleus to be a constituent part of the wall, sometimes lying enclosed between two layers of its substance, and Schwann quoted this view with seeming approval. Schwann believed, however, that in the mature cell the nucleus ceased to be functional and disappeared.

The main thesis as to the similarity of development of vegetable and animal tissues and the cellular nature of the ultimate constitution of both was supported by a mass of carefully gathered evidence which a multitude of microscopists at once confirmed, so Schwann's work became a classic almost from the moment of its publication. Of course various other workers at once disputed Schwann's claim to priority of discovery, in particular the English microscopist Valentin, who asserted, not without some show of justice, that he was working closely along the same lines. Put so, for that matter, were numerous others, as Henle, Turpin, Du-mortier, Purkinje, and Muller, all of whom Schwann himself had quoted. Moreover, there were various physiologists who earlier than any of these had foreshadowed the cell theory—notably Kaspar Friedrich Wolff, towards the close of the previous century, and Treviranus about 1807, But, as we have seen in so many other departments of science, it is one thing to foreshadow a discovery, it is quite another to give it full expression and make it germinal of other discoveries. And when Schwann put forward the explicit claim that "there is one universal principle of development for the elementary parts, of organisms, however different, and this principle is the formation of cells," he enunciated a doctrine which was for all practical purposes absolutely new and opened up a novel field for the microscopist to enter. A most important era in physiology dates from the publication of his book in 1839.

THE CELL THEORY ELABORATED

That Schwann should have gone to embryonic tissues for the establishment of his ideas was no doubt due very largely to the influence of the great Russian Karl Ernst von Baer, who about ten years earlier had published the first part of his celebrated work on embryology, and whose ideas were rapidly gaining ground, thanks largely to the advocacy of a few men, notably Johannes Muller, in Germany, and William B. Carpenter, in England, and to the fact that the improved microscope had made minute anatomy popular. Schwann's researches made it plain that the best field for the study of the animal cell is here, and a host of explorers entered the field. The result of their observations was, in the main, to confirm the claims of Schwann as to the universal prevalence of the cell. The long-current idea that animal tissues grow only as a sort of deposit from the blood-vessels was now discarded, and the fact of so-called plantlike growth of animal cells, for which Schwann contended, was universally accepted. Yet the full measure of the affinity between the two classes of cells was not for some time generally apprehended.

Indeed, since the substance that composes the cell walls of plants is manifestly very different from the limiting membrane of the animal cell, it was natural, so long as the wall was considered the most essential part of the structure, that the divergence between the two classes of cells should seem very pronounced. And for a time this was the conception of the matter that was uniformly accepted. But as time went on many observers had their attention called to the peculiar characteristics of the contents of the cell, and were led to ask themselves whether these might not be more important than had been supposed. In particular, Dr. Hugo von Mohl, professor of botany in the University of Tubingen, in the course of his exhaustive studies of the vegetable cell, was impressed with the peculiar and characteristic appearance of the cell contents. He observed universally within the cell "an opaque, viscid fluid, having granules intermingled in it," which made up the main substance of the cell, and which particularly impressed him because under certain conditions it could be seen to be actively in motion, its parts separated into filamentous streams.

Von Mohl called attention to the fact that this motion of the cell contents had been observed as long ago as 1774 by Bonaventura Corti, and rediscovered in 1807 by Treviranus, and that these observers had described the phenomenon under the "most unsuitable name of 'rotation of the cell sap.'" Von Mohl recognized that the streaming substance was something quite different from sap. He asserted that the nucleus of the cell lies within this substance and not attached to the cell wall as Schleiden had contended. He saw, too, that the chlorophyl granules, and all other of the cell contents, are incorporated with the "opaque, viscid fluid," and in 1846 he had become so impressed with the importance of this universal cell substance that he gave it the name of protoplasm. Yet in so doing he had no intention of subordinating the cell wall. The fact that Payen, in 1844, had demonstrated that the cell walls of all vegetables, high or low, are composed largely of one substance, cellulose, tended to strengthen the position of the cell wall as the really essential structure, of which the protoplasmic contents were only subsidiary products.

Meantime, however, the students of animal histology were more and more impressed with the seeming preponderance of cell contents over cell walls in the tissues they studied. They, too, found the cell to be filled with a viscid, slimy fluid capable of motion. To this Dujardin gave the name of sarcode. Presently it came to be known, through the labors of Kolliker, Nageli, Bischoff, and various others, that there are numerous lower forms of animal life which seem to be composed of this sarcode, without any cell wall whatever. The same thing seemed to be true of certain cells of higher organisms, as the blood corpuscles. Particularly in the case of cells that change their shape markedly, moving about in consequence of the streaming of their sarcode, did it seem certain that no cell wall is present, or that, if present, its role must be insignificant.

And so histologists came to question whether, after all, the cell contents rather than the enclosing wall must not be the really essential structure, and the weight of increasing observations finally left no escape from the conclusion that such is really the case. But attention being thus focalized on the cell contents, it was at once apparent that there is a far closer similarity between the ultimate particles of vegetables and those of animals than had been supposed. Cellulose and animal membrane being now regarded as more by-products, the way was clear for the recognition of the fact that vegetable protoplasm and animal sarcode are marvellously similar in appearance and general properties. The closer the observation the more striking seemed this similarity; and finally, about 1860, it was demonstrated by Heinrich de Bary and by Max Schultze that the two are to all intents and purposes identical. Even earlier Remak had reached a similar conclusion, and applied Von Mohl's word protoplasm to animal cell contents, and now this application soon became universal. Thenceforth this protoplasm was to assume the utmost importance in the physiological world, being recognized as the universal "physical basis of life," vegetable and animal alike. This amounted to the logical extension and culmination of Schwann's doctrine as to the similarity of development of the two animate kingdoms. Yet at the same time it was in effect the banishment of the cell that Schwann had defined. The word cell was retained, it is true, but it no longer signified a minute cavity. It now implied, as Schultze defined it, "a small mass of protoplasm endowed with the attributes of life." This definition was destined presently to meet with yet another modification, as we shall see; but the conception of the protoplasmic mass as the essential ultimate structure, which might or might not surround itself with a protective covering, was a permanent addition to physiological knowledge. The earlier idea had, in effect, declared the shell the most important part of the egg; this developed view assigned to the yolk its true position.

In one other important regard the theory of Schleiden and Schwann now became modified. This referred to the origin of the cell. Schwann had regarded cell growth as a kind of crystallization, beginning with the deposit of a nucleus about a granule in the intercellular substance—the cytoblastema, as Schleiden called it. But Von Mohl, as early as 1835, had called attention to the formation of new vegetable cells through the division of a pre-existing cell. Ehrenberg, another high authority of the time, contended that no such division occurs, and the matter was still in dispute when Schleiden came forward with his discovery of so-called free cell-formation within the parent cell, and this for a long time diverted attention from the process of division which Von Mohl had described. All manner of schemes of cell-formation were put forward during the ensuing years by a multitude of observers, and gained currency notwithstanding Von Mohl's reiterated contention that there are really but two ways in which the formation of new cells takes place—namely, "first, through division of older cells; secondly, through the formation of secondary cells lying free in the cavity of a cell."

But gradually the researches of such accurate observers as Unger, Nageli, Kolliker, Reichart, and Remak tended to confirm the opinion of Von Mohl that cells spring only from cells, and finally Rudolf Virchow brought the matter to demonstration about 1860. His Omnis cellula e cellula became from that time one of the accepted data of physiology. This was supplemented a little later by Fleming's Omnis nucleus e nucleo, when still more refined methods of observation had shown that the part of the cell which always first undergoes change preparatory to new cell-formation is the all-essential nucleus. Thus the nucleus was restored to the important position which Schwann and Schleiden had given it, but with greatly altered significance. Instead of being a structure generated de novo from non-cellular substance, and disappearing as soon as its function of cell-formation was accomplished, the nucleus was now known as the central and permanent feature of every cell, indestructible while the cell lives, itself the division-product of a pre-existing nucleus, and the parent, by division of its substance, of other generations of nuclei. The word cell received a final definition as "a small mass of protoplasm supplied with a nucleus."

In this widened and culminating general view of the cell theory it became clear that every animate organism, animal or vegetable, is but a cluster of nucleated cells, all of which, in each individual case, are the direct descendants of a single primordial cell of the ovum. In the developed individuals of higher organisms the successive generations of cells become marvellously diversified in form and in specific functions; there is a wonderful division of labor, special functions being chiefly relegated to definite groups of cells; but from first to last there is no function developed that is not present, in a primitive way, in every cell, however isolated; nor does the developed cell, however specialized, ever forget altogether any one of its primordial functions or capacities. All physiology, then, properly interpreted, becomes merely a study of cellular activities; and the development of the cell theory takes its place as the great central generalization in physiology of the nineteenth century. Something of the later developments of this theory we shall see in another connection.

ANIMAL CHEMISTRY

Just at the time when the microscope was opening up the paths that were to lead to the wonderful cell theory, another novel line of interrogation of the living organism was being put forward by a different set of observers. Two great schools of physiological chemistry had arisen—one under guidance of Liebig and Wohler, in Germany, the other dominated by the great French master Jean Baptiste Dumas. Liebig had at one time contemplated the study of medicine, and Dumas had achieved distinction in connection with Prevost, at Geneva, in the field of pure physiology before he turned his attention especially to chemistry. Both these masters, therefore, and Wohler as well, found absorbing interest in those phases of chemistry that have to do with the functions of living tissues; and it was largely through their efforts and the labors of their followers that the prevalent idea that vital processes are dominated by unique laws was discarded and physiology was brought within the recognized province of the chemist. So at about the time when the microscope had taught that the cell is the really essential structure of the living organism, the chemists had come to understand that every function of the organism is really the expression of a chemical change—that each cell is, in short, a miniature chemical laboratory. And it was this combined point of view of anatomist and chemist, this union of hitherto dissociated forces, that made possible the inroads into the unexplored fields of physiology that were effected towards the middle of the nineteenth century.

One of the first subjects reinvestigated and brought to proximal solution was the long-mooted question of the digestion of foods. Spallanzani and Hunter had shown in the previous century that digestion is in some sort a solution of foods; but little advance was made upon their work until 1824, when Prout detected the presence of hydrochloric acid in the gastric juice. A decade later Sprott and Boyd detected the existence of peculiar glands in the gastric mucous membrane; and Cagniard la Tour and Schwann independently discovered that the really active principle of the gastric juice is a substance which was named pepsin, and which was shown by Schwann to be active in the presence of hydrochloric acid.

Almost coincidently, in 1836, it was discovered by Purkinje and Pappenheim that another organ than the stomach—namely, the pancreas—has a share in digestion, and in the course of the ensuing decade it came to be known, through the efforts of Eberle, Valentin, and Claude Bernard, that this organ is all-important in the digestion of starchy and fatty foods. It was found, too, that the liver and the intestinal glands have each an important share in the work of preparing foods for absorption, as also has the saliva—that, in short, a coalition of forces is necessary for the digestion of all ordinary foods taken into the stomach.

And the chemists soon discovered that in each one of the essential digestive juices there is at least one substance having certain resemblances to pepsin, though acting on different kinds of food. The point of resemblance between all these essential digestive agents is that each has the remarkable property of acting on relatively enormous quantities of the substance which it can digest without itself being destroyed or apparently even altered. In virtue of this strange property, pepsin and the allied substances were spoken of as ferments, but more recently it is customary to distinguish them from such organized ferments as yeast by designating them enzymes. The isolation of these enzymes, and an appreciation of their mode of action, mark a long step towards the solution of the riddle of digestion, but it must be added that we are still quite in the dark as to the real ultimate nature of their strange activity.

In a comprehensive view, the digestive organs, taken as a whole, are a gateway between the outside world and the more intimate cells of the organism. Another equally important gateway is furnished by the lungs, and here also there was much obscurity about the exact method of functioning at the time of the revival of physiological chemistry. That oxygen is consumed and carbonic acid given off during respiration the chemists of the age of Priestley and Lavoisier had indeed made clear, but the mistaken notion prevailed that it was in the lungs themselves that the important burning of fuel occurs, of which carbonic acid is a chief product. But now that attention had been called to the importance of the ultimate cell, this misconception could not long hold its ground, and as early as 1842 Liebig, in the course of his studies of animal heat, became convinced that it is not in the lungs, but in the ultimate tissues to which they are tributary, that the true consumption of fuel takes place. Reviving Lavoisier's idea, with modifications and additions, Liebig contended, and in the face of opposition finally demonstrated, that the source of animal heat is really the consumption of the fuel taken in through the stomach and the lungs. He showed that all the activities of life are really the product of energy liberated solely through destructive processes, amounting, broadly speaking, to combustion occurring in the ultimate cells of the organism. Here is his argument:

LIEBIG ON ANIMAL HEAT

"The oxygen taken into the system is taken out again in the same forms, whether in summer or in winter; hence we expire more carbon in cold weather, and when the barometer is high, than we do in warm weather; and we must consume more or less carbon in our food in the same proportion; in Sweden more than in Sicily; and in our more temperate climate a full eighth more in winter than in summer.

"Even when we consume equal weights of food in cold and warm countries, infinite wisdom has so arranged that the articles of food in different climates are most unequal in the proportion of carbon they contain. The fruits on which the natives of the South prefer to feed do not in the fresh state contain more than twelve per cent. of carbon, while the blubber and train-oil used by the inhabitants of the arctic regions contain from sixty-six to eighty per cent. of carbon.

"It is no difficult matter, in warm climates, to study moderation in eating, and men can bear hunger for a long time under the equator; but cold and hunger united very soon exhaust the body.

"The mutual action between the elements of the food and the oxygen conveyed by the circulation of the blood to every part of the body is the source of animal heat.

"All living creatures whose existence depends on the absorption of oxygen possess within themselves a source of heat independent of surrounding objects.

"This truth applies to all animals, and extends besides to the germination of seeds, to the flowering of plants, and to the maturation of fruits. It is only in those parts of the body to which arterial blood, and with it the oxygen absorbed in respiration, is conveyed that heat is produced. Hair, wool, or feathers do not possess an elevated temperature. This high temperature of the animal body, or, as it may be called, disengagement of heat, is uniformly and under all circumstances the result of the combination of combustible substance with oxygen.

"In whatever way carbon may combine with oxygen, the act of combination cannot take place without the disengagement of heat. It is a matter of indifference whether the combination takes place rapidly or slowly, at a high or at a low temperature; the amount of heat liberated is a constant quantity. The carbon of the food, which is converted into carbonic acid within the body, must give out exactly as much heat as if it had been directly burned in the air or in oxygen gas; the only difference is that the amount of heat produced is diffused over unequal times. In oxygen the combustion is more rapid and the heat more intense; in air it is slower, the temperature is not so high, but it continues longer.

"It is obvious that the amount of heat liberated must increase or diminish with the amount of oxygen introduced in equal times by respiration. Those animals which respire frequently, and consequently consume much oxygen, possess a higher temperature than others which, with a body of equal size to be heated, take into the system less oxygen. The temperature of a child (102 degrees) is higher than that of an adult (99.5 degrees). That of birds (104 to 105.4 degrees) is higher than that of quadrupeds (98.5 to 100.4 degrees), or than that of fishes or amphibia, whose proper temperature is from 3.7 to 2.6 degrees higher than that of the medium in which they live. All animals, strictly speaking, are warm-blooded; but in those only which possess lungs is the temperature of the body independent of the surrounding medium.

"The most trustworthy observations prove that in all climates, in the temperate zones as well as at the equator or the poles, the temperature of the body in man, and of what are commonly called warm-blooded animals, is invariably the same; yet how different are the circumstances in which they live.

"The animal body is a heated mass, which bears the same relation to surrounding objects as any other heated mass. It receives heat when the surrounding objects are hotter, it loses heat when they are colder than itself. We know that the rapidity of cooling increases with the difference between the heated body and that of the surrounding medium—that is, the colder the surrounding medium the shorter the time required for the cooling of the heated body. How unequal, then, must be the loss of heat of a man at Palermo, where the actual temperature is nearly equal to that of the body, and in the polar regions, where the external temperature is from 70 to 90 degrees lower.

"Yet notwithstanding this extremely unequal loss of heat, experience has shown that the blood of an inhabitant of the arctic circle has a temperature as high as that of the native of the South, who lives in so different a medium. This fact, when its true significance is perceived, proves that the heat given off to the surrounding medium is restored within the body with great rapidity. This compensation takes place more rapidly in winter than in summer, at the pole than at the equator.

"Now in different climates the quantity of oxygen introduced into the system of respiration, as has been already shown, varies according to the temperature of the external air; the quantity of inspired oxygen increases with the loss of heat by external cooling, and the quantity of carbon or hydrogen necessary to combine with this oxygen must be increased in like ratio. It is evident that the supply of heat lost by cooling is effected by the mutual action of the elements of the food and the inspired oxygen, which combine together. To make use of a familiar, but not on that account a less just illustration, the animal body acts, in this respect, as a furnace, which we supply with fuel. It signifies nothing what intermediate forms food may assume, what changes it may undergo in the body, the last change is uniformly the conversion of carbon into carbonic acid and of its hydrogen into water; the unassimilated nitrogen of the food, along with the unburned or unoxidized carbon, is expelled in the excretions. In order to keep up in a furnace a constant temperature, we must vary the supply of fuel according to the external temperature—that is, according to the supply of oxygen.

"In the animal body the food is the fuel; with a proper supply of oxygen we obtain the heat given out during its oxidation or combustion."(3)

BLOOD CORPUSCLES, MUSCLES, AND GLANDS

Further researches showed that the carriers of oxygen, from the time of its absorption in the lungs till its liberation in the ultimate tissues, are the red corpuscles, whose function had been supposed to be the mechanical one of mixing of the blood. It transpired that the red corpuscles are composed chiefly of a substance which Kuhne first isolated in crystalline form in 1865, and which was named haemoglobin—a substance which has a marvellous affinity for oxygen, seizing on it eagerly at the lungs vet giving it up with equal readiness when coursing among the remote cells of the body. When freighted with oxygen it becomes oxyhaemoglobin and is red in color; when freed from its oxygen it takes a purple hue; hence the widely different appearance of arterial and venous blood, which so puzzled the early physiologists.

This proof of the vitally important role played by the red-blood corpuscles led, naturally, to renewed studies of these infinitesimal bodies. It was found that they may vary greatly in number at different periods in the life of the same individual, proving that they may be both developed and destroyed in the adult organism. Indeed, extended observations left no reason to doubt that the process of corpuscle formation and destruction may be a perfectly normal one—that, in short, every red-blood corpuscle runs its course and dies like any more elaborate organism. They are formed constantly in the red marrow of bones, and are destroyed in the liver, where they contribute to the formation of the coloring matter of the bile. Whether there are other seats of such manufacture and destruction of the corpuscles is not yet fully determined. Nor are histologists agreed as to whether the red-blood corpuscles themselves are to be regarded as true cells, or merely as fragments of cells budded out from a true cell for a special purpose; but in either case there is not the slightest doubt that the chief function of the red corpuscle is to carry oxygen.

If the oxygen is taken to the ultimate cells before combining with the combustibles it is to consume, it goes without saying that these combustibles themselves must be carried there also. Nor could it be in doubt that the chiefest of these ultimate tissues, as regards, quantity of fuel required, are the muscles. A general and comprehensive view of the organism includes, then, digestive apparatus and lungs as the channels of fuel-supply; blood and lymph channels as the transportation system; and muscle cells, united into muscle fibres, as the consumption furnaces, where fuel is burned and energy transformed and rendered available for the purposes of the organism, supplemented by a set of excretory organs, through which the waste products—the ashes—are eliminated from the system.

But there remain, broadly speaking, two other sets of organs whose size demonstrates their importance in the economy of the organism, yet whose functions are not accounted for in this synopsis. These are those glandlike organs, such as the spleen, which have no ducts and produce no visible secretions, and the nervous mechanism, whose central organs are the brain and spinal cord. What offices do these sets of organs perform in the great labor-specializing aggregation of cells which we call a living organism?

As regards the ductless glands, the first clew to their function was given when the great Frenchman Claude Bernard (the man of whom his admirers loved to say, "He is not a physiologist merely; he is physiology itself") discovered what is spoken of as the glycogenic function of the liver. The liver itself, indeed, is not a ductless organ, but the quantity of its biliary output seems utterly disproportionate to its enormous size, particularly when it is considered that in the case of the human species the liver contains normally about one-fifth of all the blood in the entire body. Bernard discovered that the blood undergoes a change of composition in passing through the liver. The liver cells (the peculiar forms of which had been described by Purkinje, Henle, and Dutrochet about 1838) have the power to convert certain of the substances that come to them into a starchlike compound called glycogen, and to store this substance away till it is needed by the organism. This capacity of the liver cells is quite independent of the bile-making power of the same cells; hence the discovery of this glycogenic function showed that an organ may have more than one pronounced and important specific function. But its chief importance was in giving a clew to those intermediate processes between digestion and final assimilation that are now known to be of such vital significance in the economy of the organism.

In the forty odd years that have elapsed since this pioneer observation of Bernard, numerous facts have come to light showing the extreme importance of such intermediate alterations of food-supplies in the blood as that performed by the liver. It has been shown that the pancreas, the spleen, the thyroid gland, the suprarenal capsules are absolutely essential, each in its own way, to the health of the organism, through metabolic changes which they alone seem capable of performing; and it is suspected that various other tissues, including even the muscles themselves, have somewhat similar metabolic capacities in addition to their recognized functions. But so extremely intricate is the chemistry of the substances involved that in no single case has the exact nature of the metabolisms wrought by these organs been fully made out. Each is in its way a chemical laboratory indispensable to the right conduct of the organism, but the precise nature of its operations remains inscrutable. The vast importance of the operations of these intermediate organs is unquestioned.

A consideration of the functions of that other set of organs known collectively as the nervous system is reserved for a later chapter.



VI. THEORIES OF ORGANIC EVOLUTION

GOETHE AND THE METAMORPHOSIS OF PARTS

When Coleridge said of Humphry Davy that he might have been the greatest poet of his time had he not chosen rather to be the greatest chemist, it is possible that the enthusiasm of the friend outweighed the caution of the critic. But however that may be, it is beyond dispute that the man who actually was the greatest poet of that time might easily have taken the very highest rank as a scientist had not the muse distracted his attention. Indeed, despite these distractions, Johann Wolfgang von Goethe achieved successes in the field of pure science that would insure permanent recognition for his name had he never written a stanza of poetry. Such is the versatility that marks the highest genius.

It was in 1790 that Goethe published the work that laid the foundations of his scientific reputation—the work on the Metamorphoses of Plants, in which he advanced the novel doctrine that all parts of the flower are modified or metamorphosed leaves.

"Every one who observes the growth of plants, even superficially," wrote Goethe, "will notice that certain external parts of them become transformed at times and go over into the forms of the contiguous parts, now completely, now to a greater or less degree. Thus, for example, the single flower is transformed into a double one when, instead of stamens, petals are developed, which are either exactly like the other petals of the corolla in form, and color or else still bear visible signs of their origin.

"When we observe that it is possible for a plant in this way to take a step backward, we shall give so much the more heed to the regular course of nature and learn the laws of transformation according to which she produces one part through another, and displays the most varying forms through the modification of one single organ.

"Let us first direct our attention to the plant at the moment when it develops out of the seed-kernel. The first organs of its upward growth are known by the name of cotyledons; they have also been called seed-leaves.

"They often appear shapeless, filled with new matter, and are just as thick as they are broad. Their vessels are unrecognizable and are hardly to be distinguished from the mass of the whole; they bear almost no resemblance to a leaf, and we could easily be misled into regarding them as special organs. Occasionally, however, they appear as real leaves, their vessels are capable of the most minute development, their similarity to the following leaves does not permit us to take them for special organs, but we recognize them instead to be the first leaves of the stalk.

"The cotyledons are mostly double, and there is an observation to be made here which will appear still more important as we proceed—that is, that the leaves of the first node are often paired, even when the following leaves of the stalk stand alternately upon it. Here we see an approximation and a joining of parts which nature afterwards separates and places at a distance from one another. It is still more remarkable when the cotyledons take the form of many little leaves gathered about an axis, and the stalk which grows gradually from their midst produces the following leaves arranged around it singly in a whorl. This may be observed very exactly in the growth of the pinus species. Here a corolla of needles forms at the same time a calyx, and we shall have occasion to remember the present case in connection with similar phenomena later.

"On the other hand, we observe that even the cotyledons which are most like a leaf when compared with the following leaves of the stalk are always more undeveloped or less developed. This is chiefly noticeable in their margin which is extremely simple and shows few traces of indentation.

"A few or many of the next following leaves are often already present in the seed, and lie enclosed between the cotyledons; in their folded state they are known by the name of plumules. Their form, as compared with the cotyledons and the following leaves, varies in different plants. Their chief point of variance, however, from the cotyledons is that they are flat, delicate, and formed like real leaves generally. They are wholly green, rest on a visible node, and can no longer deny their relationship to the following leaves of the stalk, to which, however, they are usually still inferior, in so far as that their margin is not completely developed.

"The further development, however, goes on ceaselessly in the leaf, from node to node; its midrib is elongated, and more or less additional ribs stretch out from this towards the sides. The leaves now appear notched, deeply indented, or composed of several small leaves, in which last case they seem to form complete little branches. The date-palm furnishes a striking example of such a successive transformation of the simplest leaf form. A midrib is elongated through a succession of several leaves, the single fan-shaped leaf becomes torn and diverted, and a very complicated leaf is developed, which rivals a branch in form.

"The transition to inflorescence takes place more or less rapidly. In the latter case we usually observe that the leaves of the stalk loose their different external divisions, and, on the other hand, spread out more or less in their lower parts where they are attached to the stalk. If the transition takes place rapidly, the stalk, suddenly become thinner and more elongated since the node of the last-developed leaf, shoots up and collects several leaves around an axis at its end.

"That the petals of the calyx are precisely the same organs which have hitherto appeared as leaves on the stalk, but now stand grouped about a common centre in an often very different form, can, as it seems to me, be most clearly demonstrated. Already in connection with the cotyledons above, we noticed a similar working of nature. The first species, while they are developing out of the seed-kernel, display a radiate crown of unmistakable needles; and in the first childhood of these plants we see already indicated that force of nature whereby when they are older their flowering and fruit-giving state will be produced.

"We see this force of nature, which collects several leaves around an axis, produce a still closer union and make these approximated, modified leaves still more unrecognizable by joining them together either wholly or partially. The bell-shaped or so-called one-petalled calices represent these cloudy connected leaves, which, being more or less indented from above, or divided, plainly show their origin.

"We can observe the transition from the calyx to the corolla in more than one instance, for, although the color of the calyx is still usually green, and like the color of the leaves of the stalk, it nevertheless often varies in one or another of its parts—at the tips, the margins, the back, or even, the inward side—while the outer still remains on green.

"The relationship of the corolla to the leaves of the stalk is shown in more than one way, since on the stalks of some plants appear leaves which are already more or less colored long before they approach inflorescence; others are fully colored when near inflorescence. Nature also goes over at once to the corolla, sometimes by skipping over the organs of the calyx, and in such a case we likewise have an opportunity to observe that leaves of the stalk become transformed into petals. Thus on the stalk of tulips, for instance, there sometimes appears an almost completely developed and colored petal. Even more remarkable is the case when such a leaf, half green and half of it belonging to the stalk, remains attached to the latter, while another colored part is raised with the corolla, and the leaf is thus torn in two.

"The relationship between the petals and stamens is very close. In some instances nature makes the transition regular—e.g., among the Canna and several plants of the same family. A true, little-modified petal is drawn together on its upper margin, and produces a pollen sac, while the rest of the petal takes the place of the stamen. In double flowers we can observe this transition in all its stages. In several kinds of roses, within the fully developed and colored petals there appear other ones which are drawn together in the middle or on the side. This drawing together is produced by a small weal, which appears as a more or less complete pollen sac, and in the same proportion the leaf approaches the simple form of a stamen.

"The pistil in many cases looks almost like a stamen without anthers, and the relationship between the formation of the two is much closer than between the other parts. In retrograde fashion nature often produces cases where the style and stigma (Narben) become retransformed into petals—that is, the Ranunculus Asiaticus becomes double by transforming the stigma and style of the fruit-receptacle into real petals, while the stamens are often found unchanged immediately behind the corolla.

"In the seed receptacles, in spite of their formation, of their special object, and of their method of being joined together, we cannot fail to recognize the leaf form. Thus, for instance, the pod would be a simple leaf folded and grown together on its margin; the siliqua would consist of more leaves folded over another; the compound receptacles would be explained as being several leaves which, being united above one centre, keep their inward parts separate and are joined on their margins. We can convince ourselves of this by actual sight when such composite capsules fall apart after becoming ripe, because then every part displays an opened pod."(1)

The theory thus elaborated of the metamorphosis of parts was presently given greater generality through extension to the animal kingdom, in the doctrine which Goethe and Oken advanced independently, that the vertebrate skull is essentially a modified and developed vertebra. These were conceptions worthy of a poet—impossible, indeed, for any mind that had not the poetic faculty of correlation. But in this case the poet's vision was prophetic of a future view of the most prosaic science. The doctrine of metamorphosis of parts soon came to be regarded as of fundamental importance.

But the doctrine had implications that few of its early advocates realized. If all the parts of a flower—sepal, petal, stamen, pistil, with their countless deviations of contour and color—are but modifications of the leaf, such modification implies a marvellous differentiation and development. To assert that a stamen is a metamorphosed leaf means, if it means anything, that in the long sweep of time the leaf has by slow or sudden gradations changed its character through successive generations, until the offspring, so to speak, of a true leaf has become a stamen. But if such a metamorphosis as this is possible—if the seemingly wide gap between leaf and stamen may be spanned by the modification of a line of organisms—where does the possibility of modification of organic type find its bounds? Why may not the modification of parts go on along devious lines until the remote descendants of an organism are utterly unlike that organism? Why may we not thus account for the development of various species of beings all sprung from one parent stock? That, too, is a poet's dream; but is it only a dream? Goethe thought not. Out of his studies of metamorphosis of parts there grew in his mind the belief that the multitudinous species of plants and animals about us have been evolved from fewer and fewer earlier parent types, like twigs of a giant tree drawing their nurture from the same primal root. It was a bold and revolutionary thought, and the world regarded it as but the vagary of a poet.

ERASMUS DARWIN

Just at the time when this thought was taking form in Goethe's brain, the same idea was germinating in the mind of another philosopher, an Englishman of international fame, Dr. Erasmus Darwin, who, while he lived, enjoyed the widest popularity as a poet, the rhymed couplets of his Botanic Garden being quoted everywhere with admiration. And posterity repudiating the verse which makes the body of the book, yet grants permanent value to the book itself, because, forsooth, its copious explanatory foot-notes furnish an outline of the status of almost every department of science of the time.

But even though he lacked the highest art of the versifier, Darwin had, beyond peradventure, the imagination of a poet coupled with profound scientific knowledge; and it was his poetic insight, correlating organisms seemingly diverse in structure and imbuing the lowliest flower with a vital personality, which led him to suspect that there are no lines of demarcation in nature. "Can it be," he queries, "that one form of organism has developed from another; that different species are really but modified descendants of one parent stock?" The alluring thought nestled in his mind and was nurtured there, and grew in a fixed belief, which was given fuller expression in his Zoonomia and in the posthumous Temple of Nature.

Here is his rendering of the idea as versified in the Temple of Nature:

"Organic life beneath the shoreless waves Was born, and nursed in Ocean's pearly caves; First forms minute, unseen by spheric glass, Move on the mud, or pierce the watery mass; These, as successive generations bloom, New powers acquire and larger limbs assume; Whence countless groups of vegetation spring, And breathing realms of fin, and feet, and wing.

"Thus the tall Oak, the giant of the wood, Which bears Britannia's thunders on the flood; The Whale, unmeasured monster of the main; The lordly lion, monarch of the plain; The eagle, soaring in the realms of air, Whose eye, undazzled, drinks the solar glare; Imperious man, who rules the bestial crowd, Of language, reason, and reflection proud, With brow erect, who scorns this earthy sod, And styles himself the image of his God— Arose from rudiments of form and sense, An embryon point or microscopic ens!"(2)

Here, clearly enough, is the idea of evolution. But in that day there was little proof forthcoming of its validity that could satisfy any one but a poet, and when Erasmus Darwin died, in 1802, the idea of transmutation of species was still but an unsubstantiated dream.

It was a dream, however, which was not confined to Goethe and Darwin. Even earlier the idea had come more or less vaguely to another great dreamer—and worker—of Germany, Immanuel Kant, and to several great Frenchmen, including De Maillet, Maupertuis, Robinet, and the famous naturalist Buffon—a man who had the imagination of a poet, though his message was couched in most artistic prose. Not long after the middle of the eighteenth century Buffon had put forward the idea of transmutation of species, and he reiterated it from time to time from then on till his death in 1788. But the time was not yet ripe for the idea of transmutation of species to burst its bonds.

And yet this idea, in a modified or undeveloped form, had taken strange hold upon the generation that was upon the scene at the close of the eighteenth century. Vast numbers of hitherto unknown species of animals had been recently discovered in previously unexplored regions of the globe, and the wise men were sorely puzzled to account for the disposal of all of these at the time of the deluge. It simplified matters greatly to suppose that many existing species had been developed since the episode of the ark by modification of the original pairs. The remoter bearings of such a theory were overlooked for the time, and the idea that American animals and birds, for example, were modified descendants of Old-World forms—the jaguar of the leopard, the puma of the lion, and so on—became a current belief with that class of humanity who accept almost any statement as true that harmonizes with their prejudices without realizing its implications.

Thus it is recorded with eclat that the discovery of the close proximity of America at the northwest with Asia removes all difficulties as to the origin of the Occidental faunas and floras, since Oriental species might easily have found their way to America on the ice, and have been modified as we find them by "the well-known influence of climate." And the persons who gave expression to this idea never dreamed of its real significance. In truth, here was the doctrine of evolution in a nutshell, and, because its ultimate bearings were not clear, it seemed the most natural of doctrines. But most of the persons who advanced it would have turned from it aghast could they have realized its import. As it was, however, only here and there a man like Buffon reasoned far enough to inquire what might be the limits of such assumed transmutation; and only here and there a Darwin or a Goethe reached the conviction that there are no limits.

LAMARCK VERSUS CUVIER

And even Goethe and Darwin had scarcely passed beyond that tentative stage of conviction in which they held the thought of transmutation of species as an ancillary belief not ready for full exposition. There was one of their contemporaries, however, who, holding the same conception, was moved to give it full explication. This was the friend and disciple of Buffon, Jean Baptiste de Lamarck. Possessed of the spirit of a poet and philosopher, this great Frenchman had also the widest range of technical knowledge, covering the entire field of animate nature. The first half of his long life was devoted chiefly to botany, in which he attained high distinction. Then, just at the beginning of the nineteenth century, he turned to zoology, in particular to the lower forms of animal life. Studying these lowly organisms, existing and fossil, he was more and more impressed with the gradations of form everywhere to be seen; the linking of diverse families through intermediate ones; and in particular with the predominance of low types of life in the earlier geological strata. Called upon constantly to classify the various forms of life in the course of his systematic writings, he found it more and more difficult to draw sharp lines of demarcation, and at last the suspicion long harbored grew into a settled conviction that there is really no such thing as a species of organism in nature; that "species" is a figment of the human imagination, whereas in nature there are only individuals.

That certain sets of individuals are more like one another than like other sets is of course patent, but this only means, said Lamarck, that these similar groups have had comparatively recent common ancestors, while dissimilar sets of beings are more remotely related in consanguinity. But trace back the lines of descent far enough, and all will culminate in one original stock. All forms of life whatsoever are modified descendants of an original organism. From lowest to highest, then, there is but one race, one species, just as all the multitudinous branches and twigs from one root are but one tree. For purposes of convenience of description, we may divide organisms into orders, families, genera, species, just as we divide a tree into root, trunk, branches, twigs, leaves; but in the one case, as in the other, the division is arbitrary and artificial.

In Philosophie Zoologique (1809), Lamarck first explicitly formulated his ideas as to the transmutation of species, though he had outlined them as early as 1801. In this memorable publication not only did he state his belief more explicitly and in fuller detail than the idea had been expressed by any predecessor, but he took another long forward step, carrying him far beyond all his forerunners except Darwin, in that he made an attempt to explain the way in which the transmutation of species had been brought about. The changes have been wrought, he said, through the unceasing efforts of each organism to meet the needs imposed upon it by its environment. Constant striving means the constant use of certain organs. Thus a bird running by the seashore is constantly tempted to wade deeper and deeper in pursuit of food; its incessant efforts tend to develop its legs, in accordance with the observed principle that the use of any organ tends to strengthen and develop it. But such slightly increased development of the legs is transmitted to the off spring of the bird, which in turn develops its already improved legs by its individual efforts, and transmits the improved tendency. Generation after generation this is repeated, until the sum of the infinitesimal variations, all in the same direction, results in the production of the long-legged wading-bird. In a similar way, through individual effort and transmitted tendency, all the diversified organs of all creatures have been developed—the fin of the fish, the wing of the bird, the hand of man; nay, more, the fish itself, the bird, the man, even. Collectively the organs make up the entire organism; and what is true of the individual organs must be true also of their ensemble, the living being.

Whatever might be thought of Lamarck's explanation of the cause of transmutation—which really was that already suggested by Erasmus Darwin—the idea of the evolution for which he contended was but the logical extension of the conception that American animals are the modified and degenerated descendants of European animals. But people as a rule are little prone to follow ideas to their logical conclusions, and in this case the conclusions were so utterly opposed to the proximal bearings of the idea that the whole thinking world repudiated them with acclaim. The very persons who had most eagerly accepted the idea of transmutation of European species into American species, and similar limited variations through changed environment, because of the relief thus given the otherwise overcrowded ark, were now foremost in denouncing such an extension of the doctrine of transmutation as Lamarck proposed.

And, for that matter, the leaders of the scientific world were equally antagonistic to the Lamarckian hypothesis. Cuvier in particular, once the pupil of Lamarck, but now his colleague, and in authority more than his peer, stood out against the transmutation doctrine with all his force. He argued for the absolute fixity of species, bringing to bear the resources of a mind which, as a mere repository of facts, perhaps never was excelled. As a final and tangible proof of his position, he brought forward the bodies of ibises that had been embalmed by the ancient Egyptians, and showed by comparison that these do not differ in the slightest particular from the ibises that visit the Nile to-day.

Cuvier's reasoning has such great historical interest—being the argument of the greatest opponent of evolution of that day—that we quote it at some length.

"The following objections," he says, "have already been started against my conclusions. Why may not the presently existing races of mammiferous land quadrupeds be mere modifications or varieties of those ancient races which we now find in the fossil state, which modifications may have been produced by change of climate and other local circumstances, and since raised to the present excessive difference by the operations of similar causes during a long period of ages?

"This objection may appear strong to those who believe in the indefinite possibility of change of form in organized bodies, and think that, during a succession of ages and by alterations of habitudes, all the species may change into one another, or one of them give birth to all the rest. Yet to these persons the following answer may be given from their own system: If the species have changed by degrees, as they assume, we ought to find traces of this gradual modification. Thus, between the palaeotherium and the species of our own day, we should be able to discover some intermediate forms; and yet no such discovery has ever been made. Since the bowels of the earth have not preserved monuments of this strange genealogy, we have no right to conclude that the ancient and now extinct species were as permanent in their forms and characters as those which exist at present; or, at least, that the catastrophe which destroyed them did not leave sufficient time for the productions of the changes that are alleged to have taken place.

"In order to reply to those naturalists who acknowledge that the varieties of animals are restrained by nature within certain limits, it would be necessary to examine how far these limits extend. This is a very curious inquiry, and in itself exceedingly interesting under a variety of relations, but has been hitherto very little attended to....

"Wild animals which subsist upon herbage feel the influence of climate a little more extensively, because there is added to it the influence of food, both in regard to its abundance and its quality. Thus the elephants of one forest are larger than those of another; their tusks also grow somewhat longer in places where their food may happen to be more favorable for the production of the substance of ivory. The same may take place in regard to the horns of stags and reindeer. But let us examine two elephants, the most dissimilar that can be conceived, we shall not discover the smallest difference in the number and articulations of the bones, the structure of the teeth, etc.........

"Nature appears also to have guarded against the alterations of species which might proceed from mixture of breeds by influencing the various species of animals with mutual aversion from one another. Hence all the cunning and all the force that man is able to exert is necessary to accomplish such unions, even between species that have the nearest resemblances. And when the mule breeds that are thus produced by these forced conjunctions happen to be fruitful, which is seldom the case, this fecundity never continues beyond a few generations, and would not probably proceed so far without a continuance of the same cares which excited it at first. Thus we never see in a wild state intermediate productions between the hare and the rabbit, between the stag and the doe, or between the marten and the weasel. But the power of man changes this established order, and continues to produce all these intermixtures of which the various species are susceptible, but which they would never produce if left to themselves.

"The degrees of these variations are proportional to the intensity of the causes that produced them—namely, the slavery or subjection under which those animals are to man. They do not proceed far in half-domesticated species. In the cat, for example, a softer or harsher fur, more brilliant or more varied colors, greater or less size—these form the whole extent of variety in the species; the skeleton of the cat of Angora differs in no regular and constant circumstances from the wild-cat of Europe...."

The most remarkable effects of the influence of man are produced upon that animal which he has reduced most completely under subjection. Dogs have been transported by mankind into every part of the world and have submitted their action to his entire direction. Regulated in their unions by the pleasure or caprice of their masters, the almost endless varieties of dogs differ from one another in color, in length, and abundance of hair, which is sometimes entirely wanting; in their natural instincts; in size, which varies in measure as one to five, mounting in some instances to more than a hundredfold in bulk; in the form of their ears, noses, and tails; in the relative length of their legs; in the progressive development of the brain, in several of the domesticated varieties occasioning alterations even in the form of the head, some of them having long, slender muzzles with a flat forehead, others having short muzzles with a forehead convex, etc., insomuch that the apparent difference between a mastiff and a water-spaniel and between a greyhound and a pugdog are even more striking than between almost any of the wild species of a genus........

It follows from these observations that animals have certain fixed and natural characters which resist the effects of every kind of influence, whether proceeding from natural causes or human interference; and we have not the smallest reason to suspect that time has any more effect on them than climate.

"I am aware that some naturalists lay prodigious stress upon the thousands which they can call into action by a dash of their pens. In such matters, however, our only way of judging as to the effects which may be produced by a long period of time is by multiplying, as it were, such as are produced by a shorter time. With this view I have endeavored to collect all the ancient documents respecting the forms of animals; and there are none equal to those furnished by the Egyptians, both in regard to their antiquity and abundance. They have not only left us representatives of animals, but even their identical bodies embalmed and preserved in the catacombs.

"I have examined, with the greatest attention, the engraved figures of quadrupeds and birds brought from Egypt to ancient Rome, and all these figures, one with another, have a perfect resemblance to their intended objects, such as they still are to-day.

"From all these established facts, there does not seem to be the smallest foundation for supposing that the new genera which I have discovered or established among extraneous fossils, such as the paleoetherium, anoplotherium, megalonyx, mastodon, pterodactylis, etc., have ever been the sources of any of our present animals, which only differ so far as they are influenced by time or climate. Even if it should prove true, which I am far from believing to be the case, that the fossil elephants, rhinoceroses, elks, and bears do not differ further from the existing species of the same genera than the present races of dogs differ among themselves, this would by no means be a sufficient reason to conclude that they were of the same species; since the races or varieties of dogs have been influenced by the trammels of domesticity, which those other animals never did, and indeed never could, experience."(3)

To Cuvier's argument from the fixity of Egyptian mummified birds and animals, as above stated, Lamarck replied that this proved nothing except that the ibis had become perfectly adapted to its Egyptian surroundings in an early day, historically speaking, and that the climatic and other conditions of the Nile Valley had not since then changed. His theory, he alleged, provided for the stability of species under fixed conditions quite as well as for transmutation under varying conditions.

But, needless to say, the popular verdict lay with Cuvier; talent won for the time against genius, and Lamarck was looked upon as an impious visionary. His faith never wavered, however. He believed that he had gained a true insight into the processes of animate nature, and he reiterated his hypotheses over and over, particularly in the introduction to his Histoire Naturelle des Animaux sans Vertebres, in 1815, and in his Systeme des Connaissances Positives de l'Homme, in 1820. He lived on till 1829, respected as a naturalist, but almost unrecognized as a prophet.

TENTATIVE ADVANCES

While the names of Darwin and Goethe, and in particular that of Lamarck, must always stand out in high relief in this generation as the exponents of the idea of transmutation of species, there are a few others which must not be altogether overlooked in this connection. Of these the most conspicuous is that of Gottfried Reinhold Treviranus, a German naturalist physician, professor of mathematics in the lyceum at Bremen.

It was an interesting coincidence that Treviranus should have published the first volume of his Biologie, oder Philosophie der lebenden Natur, in which his views on the transmutation of species were expounded, in 1802, the same twelvemonth in which Lamarck's first exposition of the same doctrine appeared in his Recherches sur l'Organisation des Corps Vivants. It is singular, too, that Lamarck, in his Hydrogelogie of the same date, should independently have suggested "biology" as an appropriate word to express the general science of living things. It is significant of the tendency of thought of the time that the need of such a unifying word should have presented itself simultaneously to independent thinkers in different countries.

That same memorable year, Lorenz Oken, another philosophical naturalist, professor in the University of Zurich, published the preliminary outlines of his Philosophie der Natur, which, as developed through later publications, outlined a theory of spontaneous generation and of evolution of species. Thus it appears that this idea was germinating in the minds of several of the ablest men of the time during the first decade of our century. But the singular result of their various explications was to give sudden check to that undercurrent of thought which for some time had been setting towards this conception. As soon as it was made clear whither the concession that animals may be changed by their environment must logically trend, the recoil from the idea was instantaneous and fervid. Then for a generation Cuvier was almost absolutely dominant, and his verdict was generally considered final.

There was, indeed, one naturalist of authority in France who had the hardihood to stand out against Cuvier and his school, and who was in a position to gain a hearing, though by no means to divide the following. This was Etienne Geoffroy Saint-Hilaire, the famous author of the Philosophie Anatomique, and for many years the colleague of Lamarck at the Jardin des Plantes. Like Goethe, Geoffroy was pre-eminently an anatomist, and, like the great German, he had early been impressed with the resemblances between the analogous organs of different classes of beings. He conceived the idea that an absolute unity of type prevails throughout organic nature as regards each set of organs. Out of this idea grew his gradually formed belief that similarity of structure might imply identity of origin—that, in short, one species of animal might have developed from another.

Geoffroy's grasp of this idea of transmutation was by no means so complete as that of Lamarck, and he seems never to have fully determined in his own mind just what might be the limits of such development of species. Certainly he nowhere includes all organic creatures in one line of descent, as Lamarck had done; nevertheless, he held tenaciously to the truth as he saw it, in open opposition to Cuvier, with whom he held a memorable debate at the Academy of Sciences in 1830—the debate which so aroused the interest and enthusiasm of Goethe, but which, in the opinion of nearly every one else, resulted in crushing defeat for Geoffrey, and brilliant, seemingly final, victory for the advocate of special creation and the fixity of species.

With that all ardent controversy over the subject seemed to end, and for just a quarter of a century to come there was published but a single argument for transmutation of species which attracted any general attention whatever. This oasis in a desert generation was a little book called Vestiges of the Natural History of Creation, which appeared anonymously in England in 1844, and which passed through numerous editions, and was the subject of no end of abusive and derisive comment. This book, the authorship of which remained for forty years a secret, is now conceded to have been the work of Robert Chambers, the well-known English author and publisher. The book itself is remarkable as being an avowed and unequivocal exposition of a general doctrine of evolution, its view being as radical and comprehensive as that of Lamarck himself. But it was a resume of earlier efforts rather than a new departure, to say nothing of its technical shortcomings, which may best be illustrated by a quotation.

"The whole question," says Chambers, "stands thus: For the theory of universal order—that is, order as presiding in both the origin and administration of the world—we have the testimony of a vast number of facts in nature, and this one in addition—that whatever is left from the domain of ignorance, and made undoubted matter of science, forms a new support to the same doctrine. The opposite view, once predominant, has been shrinking for ages into lesser space, and now maintains a footing only in a few departments of nature which happen to be less liable than others to a clear investigation. The chief of these, if not almost the only one, is the origin of the organic kingdoms. So long as this remains obscure, the supernatural will have a certain hold upon enlightened persons. Should it ever be cleared up in a way that leaves no doubt of a natural origin of plants and animals, there must be a complete revolution in the view which is generally taken of the relation of the Father of our being.

"This prepares the way for a few remarks on the present state of opinion with regard to the origin of organic nature. The great difficulty here is the apparent determinateness of species. These forms of life being apparently unchangeable, or at least always showing a tendency to return to the character from which they have diverged, the idea arises that there can have been no progression from one to another; each must have taken its special form, independently of other forms, directly from the appointment of the Creator. The Edinburgh Review writer says, 'they were created by the hand of God and adapted to the conditions of the period.' Now it is, in the first place, not certain that species constantly maintain a fixed character, for we have seen that what were long considered as determinate species have been transmuted into others. Passing, however, from this fact, as it is not generally received among men of science, there remain some great difficulties in connection with the idea of special creation. First we should have to suppose, as pointed out in my former volume, a most startling diversity of plan in the divine workings, a great general plan or system of law in the leading events of world-making, and a plan of minute, nice operation, and special attention in some of the mere details of the process. The discrepancy between the two conceptions is surely overpowering, when we allow ourselves to see the whole matter in a steady and rational light. There is, also, the striking fact of an ascertained historical progress of plants and animals in the order of their organization; marine and cellular plants and invertebrated animals first, afterwards higher examples of both. In an arbitrary system we had surely no reason to expect mammals after reptiles; yet in this order they came. The writer in the Edinburgh Review speaks of animals as coming in adaptation to conditions, but this is only true in a limited sense. The groves which formed the coal-beds might have been a fitting habitation for reptiles, birds, and mammals, as such groves are at the present day; yet we see none of the last of these classes and hardly any traces of the two first at that period of the earth. Where the iguanodon lived the elephant might have lived, but there was no elephant at that time. The sea of the Lower Silurian era was capable of supporting fish, but no fish existed. It hence forcibly appears that theatres of life must have remained unserviceable, or in the possession of a tenantry inferior to what might have enjoyed them, for many ages: there surely would have been no such waste allowed in a system where Omnipotence was working upon the plan of minute attention to specialities. The fact seems to denote that the actual procedure of the peopling of the earth was one of a natural kind, requiring a long space of time for its evolution. In this supposition the long existence of land without land animals, and more particularly without the noblest classes and orders, is only analogous to the fact, not nearly enough present to the minds of a civilized people, that to this day the bulk of the earth is a waste as far as man is concerned.

"Another startling objection is in the infinite local variation of organic forms. Did the vegetable and animal kingdoms consist of a definite number of species adapted to peculiarities of soil and climate, and universally distributed, the fact would be in harmony with the idea of special exertion. But the truth is that various regions exhibit variations altogether without apparent end or purpose. Professor Henslow enumerates forty-five distinct flowers or sets of plants upon the surface of the earth, notwithstanding that many of these would be equally suitable elsewhere. The animals of different continents are equally various, few species being the same in any two, though the general character may conform. The inference at present drawn from this fact is that there must have been, to use the language of the Rev. Dr. Pye Smith, 'separate and original creations, perhaps at different and respectively distinct epochs.' It seems hardly conceivable that rational men should give an adherence to such a doctrine when we think of what it involves. In the single fact that it necessitates a special fiat of the inconceivable Author of this sand-cloud of worlds to produce the flora of St. Helena, we read its more than sufficient condemnation. It surely harmonizes far better with our general ideas of nature to suppose that, just as all else in this far-spread science was formed on the laws impressed upon it at first by its Author, so also was this. An exception presented to us in such a light appears admissible only when we succeed in forbidding our minds to follow out those reasoning processes to which, by another law of the Almighty, they tend, and for which they are adapted."(4)

Such reasoning as this naturally aroused bitter animadversions, and cannot have been without effect in creating an undercurrent of thought in opposition to the main trend of opinion of the time. But the book can hardly be said to have done more than that. Indeed, some critics have denied it even this merit. After its publication, as before, the conception of transmutation of species remained in the popular estimation, both lay and scientific, an almost forgotten "heresy."

It is true that here and there a scientist of greater or less repute—as Von Buch, Meckel, and Von Baer in Germany, Bory Saint-Vincent in France, Wells, Grant, and Matthew in England, and Leidy in America—had expressed more or less tentative dissent from the doctrine of special creation and immutability of species, but their unaggressive suggestions, usually put forward in obscure publications, and incidentally, were utterly overlooked and ignored. And so, despite the scientific advances along many lines at the middle of the century, the idea of the transmutability of organic races had no such prominence, either in scientific or unscientific circles, as it had acquired fifty years before. Special creation held the day, seemingly unopposed.

DARWIN AND THE ORIGIN OF SPECIES

But even at this time the fancied security of the special-creation hypothesis was by no means real. Though it seemed so invincible, its real position was that of an apparently impregnable fortress beneath which, all unbeknown to the garrison, a powder-mine has been dug and lies ready for explosion. For already there existed in the secluded work-room of an English naturalist, a manuscript volume and a portfolio of notes which might have sufficed, if given publicity, to shatter the entire structure of the special-creation hypothesis. The naturalist who, by dint of long and patient effort, had constructed this powder-mine of facts was Charles Robert Darwin, grandson of the author of Zoonomia.

As long ago as July 1, 1837, young Darwin, then twenty-eight years of age, had opened a private journal, in which he purposed to record all facts that came to him which seemed to have any bearing on the moot point of the doctrine of transmutation of species. Four or five years earlier, during the course of that famous trip around the world with Admiral Fitzroy, as naturalist to the Beagle, Darwin had made the personal observations which first tended to shake his belief of the fixity of species. In South America, in the Pampean formation, he had discovered "great fossil animals covered with armor like that on the existing armadillos," and had been struck with this similarity of type between ancient and existing faunas of the same region. He was also greatly impressed by the manner in which closely related species of animals were observed to replace one another as he proceeded southward over the continent; and "by the South-American character of most of the productions of the Galapagos Archipelago, and more especially by the manner in which they differ slightly on each island of the group, none of the islands appearing to be very ancient in a geological sense."

At first the full force of these observations did not strike him; for, under sway of Lyell's geological conceptions, he tentatively explained the relative absence of life on one of the Galapagos Islands by suggesting that perhaps no species had been created since that island arose. But gradually it dawned upon him that such facts as he had observed "could only be explained on the supposition that species gradually become modified." From then on, as he afterwards asserted, the subject haunted him; hence the journal of 1837.

It will thus be seen that the idea of the variability of species came to Charles Darwin as an inference from personal observations in the field, not as a thought borrowed from books. He had, of course, read the works of his grandfather much earlier in life, but the arguments of Zoonomia and The Temple of Nature had not served in the least to weaken his acceptance of the current belief in fixity of species. Nor had he been more impressed with the doctrine of Lamarck, so closely similar to that of his grandfather. Indeed, even after his South-American experience had aroused him to a new point of view he was still unable to see anything of value in these earlier attempts at an explanation of the variation of species. In opening his journal, therefore, he had no preconceived notion of upholding the views of these or any other makers of hypotheses, nor at the time had he formulated any hypothesis of his own. His mind was open and receptive; he was eager only for facts which might lead him to an understanding of a problem which seemed utterly obscure. It was something to feel sure that species have varied; but how have such variations been brought about?

It was not long before Darwin found a clew which he thought might lead to the answer he sought. In casting about for facts he had soon discovered that the most available field for observation lay among domesticated animals, whose numerous variations within specific lines are familiar to every one. Thus under domestication creatures so tangibly different as a mastiff and a terrier have sprung from a common stock. So have the Shetland pony, the thoroughbred, and the draught-horse. In short, there is no domesticated animal that has not developed varieties deviating more or less widely from the parent stock. Now, how has this been accomplished? Why, clearly, by the preservation, through selective breeding, of seemingly accidental variations. Thus one horseman, by constantly selecting animals that "chance" to have the right build and stamina, finally develops a race of running-horses; while another horseman, by selecting a different series of progenitors, has developed a race of slow, heavy draught animals.

So far, so good; the preservation of "accidental" variations through selective breeding is plainly a means by which races may be developed that are very different from their original parent form. But this is under man's supervision and direction. By what process could such selection be brought about among creatures in a state of nature? Here surely was a puzzle, and one that must be solved before another step could be taken in this direction.

The key to the solution of this puzzle came into Darwin's mind through a chance reading of the famous essay on "Population" which Thomas Robert Malthus had published almost half a century before. This essay, expositing ideas by no means exclusively original with Malthus, emphasizes the fact that organisms tend to increase at a geometrical ratio through successive generations, and hence would overpopulate the earth if not somehow kept in check. Cogitating this thought, Darwin gained a new insight into the processes of nature. He saw that in virtue of this tendency of each race of beings to overpopulate the earth, the entire organic world, animal and vegetable, must be in a state of perpetual carnage and strife, individual against individual, fighting for sustenance and life.

That idea fully imagined, it becomes plain that a selective influence is all the time at work in nature, since only a few individuals, relatively, of each generation can come to maturity, and these few must, naturally, be those best fitted to battle with the particular circumstances in the midst of which they are placed. In other words, the individuals best adapted to their surroundings will, on the average, be those that grow to maturity and produce offspring. To these offspring will be transmitted the favorable peculiarities. Thus these peculiarities will become permanent, and nature will have accomplished precisely what the human breeder is seen to accomplish. Grant that organisms in a state of nature vary, however slightly, one from another (which is indubitable), and that such variations will be transmitted by a parent to its offspring (which no one then doubted); grant, further, that there is incessant strife among the various organisms, so that only a small proportion can come to maturity—grant these things, said Darwin, and we have an explanation of the preservation of variations which leads on to the transmutation of species themselves.

This wonderful coign of vantage Darwin had reached by 1839. Here was the full outline of his theory; here were the ideas which afterwards came to be embalmed in familiar speech in the phrases "spontaneous variation," and the "survival of the fittest," through "natural selection." After such a discovery any ordinary man would at once have run through the streets of science, so to speak, screaming "Eureka!" Not so Darwin. He placed the manuscript outline of his theory in his portfolio, and went on gathering facts bearing on his discovery. In 1844 he made an abstract in a manuscript book of the mass of facts by that time accumulated. He showed it to his friend Hooker, made careful provision for its publication in the event of his sudden death, then stored it away in his desk and went ahead with the gathering of more data. This was the unexploded powder-mine to which I have just referred.

Twelve years more elapsed—years during which the silent worker gathered a prodigious mass of facts, answered a multitude of objections that arose in his own mind, vastly fortified his theory. All this time the toiler was an invalid, never knowing a day free from illness and discomfort, obliged to husband his strength, never able to work more than an hour and a half at a stretch; yet he accomplished what would have been vast achievements for half a dozen men of robust health. Two friends among the eminent scientists of the day knew of his labors—Sir Joseph Hooker, the botanist, and Sir Charles Lyell, the geologist. Gradually Hooker had come to be more than half a convert to Darwin's views. Lyell was still sceptical, yet he urged Darwin to publish his theory without further delay lest he be forestalled. At last the patient worker decided to comply with this advice, and in 1856 he set to work to make another and fuller abstract of the mass of data he had gathered.

And then a strange thing happened. After Darwin had been at work on his "abstract" about two years, but before he had published a line of it, there came to him one day a paper in manuscript, sent for his approval by a naturalist friend named Alfred Russel Wallace, who had been for some time at work in the East India Archipelago. He read the paper, and, to his amazement, found that it contained an outline of the same theory of "natural selection" which he himself had originated and for twenty years had worked upon. Working independently, on opposite sides of the globe, Darwin and Wallace had hit upon the same explanation of the cause of transmutation of species. "Were Wallace's paper an abstract of my unpublished manuscript of 1844," said Darwin, "it could not better express my ideas."

Here was a dilemma. To publish this paper with no word from Darwin would give Wallace priority, and wrest from Darwin the credit of a discovery which he had made years before his codiscoverer entered the field. Yet, on the other hand, could Darwin honorably do otherwise than publish his friend's paper and himself remain silent? It was a complication well calculated to try a man's soul. Darwin's was equal to the test. Keenly alive to the delicacy of the position, he placed the whole matter before his friends Hooker and Lyell, and left the decision as to a course of action absolutely to them. Needless to say, these great men did the one thing which insured full justice to all concerned. They counselled a joint publication, to include on the one hand Wallace's paper, and on the other an abstract of Darwin's ideas, in the exact form in which it had been outlined by the author in a letter to Asa Gray in the previous year—an abstract which was in Gray's hands before Wallace's paper was in existence. This joint production, together with a full statement of the facts of the case, was presented to the Linnaean Society of London by Hooker and Lyell on the evening of July 1, 1858, this being, by an odd coincidence, the twenty-first anniversary of the day on which Darwin had opened his journal to collect facts bearing on the "species question." Not often before in the history of science has it happened that a great theory has been nurtured in its author's brain through infancy and adolescence to its full legal majority before being sent out into the world.

Thus the fuse that led to the great powder-mine had been lighted. The explosion itself came more than a year later, in November, 1859, when Darwin, after thirteen months of further effort, completed the outline of his theory, which was at first begun as an abstract for the Linnaean Society, but which grew to the size of an independent volume despite his efforts at condensation, and which was given that ever-to-be-famous title, The Origin of Species by Means of Natural Selection, or the Preservation of Favored Races in the Struggle for Life. And what an explosion it was! The joint paper of 1858 had made a momentary flare, causing the hearers, as Hooker said, to "speak of it with bated breath," but beyond that it made no sensation. What the result was when the Origin itself appeared no one of our generation need be told. The rumble and roar that it made in the intellectual world have not yet altogether ceased to echo after more than forty years of reverberation.

NEW CHAMPIONS

To the Origin of Species, then, and to its author, Charles Darwin, must always be ascribed chief credit for that vast revolution in the fundamental beliefs of our race which has come about since 1859, and which made the second half of the century memorable. But it must not be overlooked that no such sudden metamorphosis could have been effected had it not been for the aid of a few notable lieutenants, who rallied to the standards of the leader immediately after the publication of the Origin. Darwin had all along felt the utmost confidence in the ultimate triumph of his ideas. "Our posterity," he declared, in a letter to Hooker, "will marvel as much about the current belief (in special creation) as we do about fossil shells having been thought to be created as we now see them." But he fully realized that for the present success of his theory of transmutation the championship of a few leaders of science was all-essential. He felt that if he could make converts of Hooker and Lyell and of Thomas Henry Huxley at once, all would be well.

His success in this regard, as in others, exceeded his expectations. Hooker was an ardent disciple from reading the proof-sheets before the book was published; Lyell renounced his former beliefs and fell into line a few months later; while Huxley, so soon as he had mastered the central idea of natural selection, marvelled that so simple yet all-potent a thought had escaped him so long, and then rushed eagerly into the fray, wielding the keenest dialectic blade that was drawn during the entire controversy. Then, too, unexpected recruits were found in Sir John Lubbock and John Tyndall, who carried the war eagerly into their respective territories; while Herbert Spencer, who had advocated a doctrine of transmutation on philosophic grounds some years before Darwin published the key to the mystery—and who himself had barely escaped independent discovery of that key—lent his masterful influence to the cause. In America the famous botanist Asa Gray, who had long been a correspondent of Darwin's but whose advocacy of the new theory had not been anticipated, became an ardent propagandist; while in Germany Ernst Heinrich Haeckel, the youthful but already noted zoologist, took up the fight with equal enthusiasm.

Against these few doughty champions—with here and there another of less general renown—was arrayed, at the outset, practically all Christendom. The interest of the question came home to every person of intelligence, whatever his calling, and the more deeply as it became more and more clear how far-reaching are the real bearings of the doctrine of natural selection. Soon it was seen that should the doctrine of the survival of the favored races through the struggle for existence win, there must come with it as radical a change in man's estimate of his own position as had come in the day when, through the efforts of Copernicus and Galileo, the world was dethroned from its supposed central position in the universe. The whole conservative majority of mankind recoiled from this necessity with horror. And this conservative majority included not laymen merely, but a vast preponderance of the leaders of science also.

With the open-minded minority, on the other hand, the theory of natural selection made its way by leaps and bounds. Its delightful simplicity—which at first sight made it seem neither new nor important—coupled with the marvellous comprehensiveness of its implications, gave it a hold on the imagination, and secured it a hearing where other theories of transmutation of species had been utterly scorned. Men who had found Lamarck's conception of change through voluntary effort ridiculous, and the vaporings of the Vestiges altogether despicable, men whose scientific cautions held them back from Spencer's deductive argument, took eager hold of that tangible, ever-present principle of natural selection, and were led on and on to its goal. Hour by hour the attitude of the thinking world towards this new principle changed; never before was so great a revolution wrought so suddenly.

Nor was this merely because "the times were ripe" or "men's minds prepared for evolution." Darwin himself bears witness that this was not altogether so. All through the years in which he brooded this theory he sounded his scientific friends, and could find among them not one who acknowledged a doctrine of transmutation. The reaction from the stand-point of Lamarck and Erasmus Darwin and Goethe had been complete, and when Charles Darwin avowed his own conviction he expected always to have it met with ridicule or contempt. In 1857 there was but one man speaking with any large degree of authority in the world who openly avowed a belief in transmutation of species—that man being Herbert Spencer. But the Origin of Species came, as Huxley has said, like a flash in the darkness, enabling the benighted voyager to see the way. The score of years during which its author had waited and worked had been years well spent. Darwin had become, as he himself says, a veritable Croesus, "overwhelmed with his riches in facts"—facts of zoology, of selective artificial breeding, of geographical distribution of animals, of embryology, of paleontology. He had massed his facts about his theory, condensed them and recondensed, until his volume of five hundred pages was an encyclopaedia in scope. During those long years of musing he had thought out almost every conceivable objection to his theory, and in his book every such objection was stated with fullest force and candor, together with such reply as the facts at command might dictate. It was the force of those twenty years of effort of a master-mind that made the sudden breach in the breaswtork{sic} of current thought.

Once this breach was effected the work of conquest went rapidly on. Day by day squads of the enemy capitulated and struck their arms. By the time another score of years had passed the doctrine of evolution had become the working hypothesis of the scientific world. The revolution had been effected.

And from amid the wreckage of opinion and belief stands forth the figure of Charles Darwin, calm, imperturbable, serene; scatheless to ridicule, contumely, abuse; unspoiled by ultimate success; unsullied alike by the strife and the victory—take him for all in all, for character, for intellect, for what he was and what he did, perhaps the most Socratic figure of the century. When, in 1882, he died, friend and foe alike conceded that one of the greatest sons of men had rested from his labors, and all the world felt it fitting that the remains of Charles Darwin should be entombed in Westminster Abbey close beside the honored grave of Isaac Newton. Nor were there many who would dispute the justice of Huxley's estimate of his accomplishment: "He found a great truth trodden under foot. Reviled by bigots, and ridiculed by all the world, he lived long enough to see it, chiefly by his own efforts, irrefragably established in science, inseparably incorporated with the common thoughts of men, and only hated and feared by those who would revile but dare not."

THE ORIGIN OF THE FITTEST

Wide as are the implications of the great truth which Darwin and his co-workers established, however, it leaves quite untouched the problem of the origin of those "favored variations" upon which it operates. That such variations are due to fixed and determinate causes no one understood better than Darwin; but in his original exposition of his doctrine he made no assumption as to what these causes are. He accepted the observed fact of variation—as constantly witnessed, for example, in the differences between parents and offspring—and went ahead from this assumption.

But as soon as the validity of the principle of natural selection came to be acknowledged speculators began to search for the explanation of those variations which, for purposes of argument, had been provisionally called "spontaneous." Herbert Spencer had all along dwelt on this phase of the subject, expounding the Lamarckian conceptions of the direct influence of the environment (an idea which had especially appealed to Buffon and to Geoffroy Saint-Hilaire), and of effort in response to environment and stimulus as modifying the individual organism, and thus supplying the basis for the operation of natural selection. Haeckel also became an advocate of this idea, and presently there arose a so-called school of neo-Lamarckians, which developed particular strength and prominence in America under the leadership of Professors A. Hyatt and E. D. Cope.

But just as the tide of opinion was turning strongly in this direction, an utterly unexpected obstacle appeared in the form of the theory of Professor August Weismann, put forward in 1883, which antagonized the Lamarckian conception (though not touching the Darwinian, of which Weismann is a firm upholder) by denying that individual variations, however acquired by the mature organism, are transmissible. The flurry which this denial created has not yet altogether subsided, but subsequent observations seem to show that it was quite disproportionate to the real merits of the case. Notwithstanding Professor Weismann's objections, the balance of evidence appears to favor the view that the Lamarckian factor of acquired variations stands as the complement of the Darwinian factor of natural selection in effecting the transmutation of species.