PART II.—Do Species wear out, and, if not, why not?—Implication of the Darwinian Theory that Species are unlimited in Existence.—Examination of an Opposite Doctrine maintained by Naudin.—Evidence that Species may die out from Inherent Causes only indirect and inferential from Arrangements to secure Wide Breeding—Physiological Import of Sexes—Doubtful whether Sexual Reproduction with Wide Breeding is a Preventive or only a Palliative of Decrepitude in Species.— Darwinian Hypothesis must suppose the Former (continued 6)

The complete oblivion into which this paper and the whole subject had fallen is the more remarkable when it is seen that both are briefly but explicitly referred to in Elliott's book, with which botanists are familiar.

It is not so wonderful that the far earlier allusion to these facts by the younger Bartram should have been overlooked or disregarded. With the genuine love of Nature and fondness for exploration, 'William Bartram did not inherit the simplicity of his father, the earliest native botanist of this country. Fine writing was his foible; and the preface to his well-known "Travels" (published at Philadelphia in 1791) is its full-blown illustration, sometimes perhaps deserving the epithet which he applies to the palms of Florida—that of pomposity. In this preface he declares that "all the Sarracenias are insect-catchers, and so is the Drosera rotundifolia. Whether the insects caught in their leaves, and which dissolve and mix with the fluid, serve for aliment or support to these kind of plants is doubtful," he thinks, but he should be credited with the suggestion. In one sentence he speaks of the quantities of insects which, "being invited down to sip the mellifluous exuvia from the interior surface of the tube, where they inevitably perish," being prevented from returning by the stiff hairs all pointing downward. This, if it refers to the sweet secretion, would place it below, and not, as it is, above the bristly surface, while the liquid below, charged with decomposing insects, is declared in an earlier sentence to be "cool and animating, limpid as the morning dew." Bartram was evidently writing from memory; and it is very doubtful if he ever distinctly recognized the sweet exudation which entices insects.

Why should these plants take to organic food more than others? If we cannot answer the question, we may make a probable step toward it. For plants that are not parasitic, these, especially the sundews, have much less than the ordinary amount of chlorophyll—that is, of the universal leaf-green upon which the formation of organic matter out of inorganic materials depends. These take it instead of making it, to a certain extent.

What is the bearing of these remarkable adaptations and operations upon doctrines of evolution? There seems here to be a field on which the specific creationist, the evolutionist with design, and the necessary evolutionist, may fight out an interesting, if not decisive, "triangular duel."

XI

INSECTIVOROUS AND

CLIMBING PLANTS [XI-1]

(The Nation, January 6 and 13, 1876)

"Minerals grow; vegetables grow and live; animals grow, live, and feel;" this is the well-worn, not to say out-worn, diagnosis of the three kingdoms by Linnaeus. It must be said of it that the agreement indicated in the first couplet is unreal, and that the distinction declared in the second is evanescent. Crystals do not grow at all in the sense that plants and animals grow. On the other hand, if a response to external impressions by special movements is evidence of feeling, vegetables share this endowment with animals; while, if conscious feeling is meant, this can be affirmed only of the higher animals. What appears to remain true is, that the difference is one of successive addition. That the increment in the organic world is of many steps; that in the long series no absolute lines separate, or have always separated, organisms which barely respond to impressions from those which more actively and variously respond, and even from those that consciously so respond—this, as we all know, is what the author of the works before us has undertaken to demonstrate. Without reference here either to that part of the series with which man is connected, and in some sense or other forms a part of, or to that lower limbo where the two organic kingdoms apparently merge—or whence, in evolutionary phrase, they have emerged—Mr. Darwin, in the present volumes, directs our attention to the behavior of the highest plants alone. He shows that some (and he might add that all) of them execute movements for their own advantage, and that some capture and digest living prey. When plants are seen to move and to devour, what faculties are left that are distinctively animal?

As to insectivorous or otherwise carnivorous plants, we have so recently here discussed this subject—before it attained to all this new popularity—that a brief account of Mr. Darwin's investigation may suffice.[XI-2] It is full of interest as a physiological research, and is a model of its kind, as well for the simplicity and directness of the means employed as for the clearness with which the results are brought out—results which any one may verify now that the way to them is pointed out, and which, surprising as they are, lose half their wonder in the ease and sureness with which they seem to have been reached.

Rather more than half the volume is devoted to one subject, the round-leaved sundew (Drosera rotundifolia), a rather common plant in the northern temperate zone. That flies stick fast to its leaves, being limed by the tenacious seeming dew-drops which stud its upper face and margins, had long been noticed in Europe and in this country. We have heard hunters and explorers in our Northern woods refer with satisfaction to the fate which in this way often befalls one of their plagues, the black fly of early summer. And it was known to some observant botanists in the last century, although forgotten or discredited in this, that an insect caught on the viscid glands it has happened to alight upon is soon fixed by many more—not merely in consequence of its struggles, but by the spontaneous incurvation of the stalks of surrounding and untouched glands; and even the body of the leaf had been observed to incurve or become cup-shaped so as partly to involve the captive insect.

Mr. Darwin's peculiar investigations not only confirm all this, but add greater wonders. They relate to the sensitiveness of these tentacles, as he prefers to call them, and the mode in which it is manifested; their power of absorption; their astonishing discernment of the presence of animal or other soluble azotized matter, even in quantities so minute as to rival the spectroscope—that most exquisite instrument of modern research—in delicacy; and, finally, they establish the fact of a true digestion, in all essential respects similar to that of the stomach of animals.

First as to sensitiveness and movement. Sensitiveness is manifested by movement or change of form in response to an external impression. The sensitiveness in the sundew is all in the gland which surmounts the tentacle. To incite movement or other action, it is necessary that the gland itself should be reached. Anything laid on the surface of the viscid drop, the spherule of clear, glairy liquid which it secretes, produces no effect unless it sinks through to the gland; or unless the substance is soluble and reaches it in solution, which, in the case of certain substances, has the same effect. But the glands themselves do not move, nor does any neighboring portion of the tentacle. The outer and longer tentacles bend inward (toward the centre of the leaf) promptly, when the gland is irritated or stimulated, sweeping through an arc of 1800 or less, or more—the quickness and the extent of the inflection depending, in equally vigorous leaves, upon the amount of irritation or stimulation, and also upon its kind. A tentacle with a particle of raw meat on its gland sometimes visibly begins to bend in ten seconds, becomes strongly incurved in five minutes, and its tip reaches the centre of the leaf in half an hour; but this is a case of extreme rapidity. A particle of cinder, chalk, or sand, will also incite the bending, if actually brought in contact with the gland, not merely resting on the drop; but the inflection is then much less pronounced and more transient. Even a bit of thin human hair, only 1/8000 of an inch in length, weighing only the 1/78740 of a grain, and largely supported by the viscid secretion, suffices to induce movement; but, on the other hand, one or two momentary, although rude, touches with a hard object produce no effect, although a repeated touch or the slightest pressure, such as that of a gnat's foot, prolonged for a short time, causes bending. The seat of the movement is wholly or nearly confined to a portion of the lower part of the tentacle, above the base, where local irritation produces not the slightest effect. The movement takes place only in response to some impression made upon its own gland at the distant extremity, or upon other glands far more remote. For if one of these members suffers irritation the others sympathize with it. Very noteworthy is the correlation between the central tentacles, upon which an insect is most likely to alight, and these external and larger ones, which, in proportion to their distance from the centre, take the larger share in the movement. The shorter central ones do not move at all when a bit of meat, or a crushed fly, or a particle of a salt of ammonia, or the like, is placed upon them; but they transmit their excitation across the leaf to the surrounding tentacles on all sides; and they, although absolutely untouched, as they successively receive the mysterious impulse, bend strongly inward, just as they do when their own glands are excited. Whenever a tentacle bends in obedience to an impulse from its own gland, the movement is always toward the centre of the leaf; and this also takes place, as we have seen, when an exciting object is lodged at the centre. But when the object is placed upon either half of the leaf, the impulse radiating thence causes all the surrounding untouched tentacles to bend with precision toward the point of excitement, even the central tentacles, which are motionless when themselves charged, now responding to the call. The inflection which follows mechanical irritation or the presence of any inorganic or insoluble body is transient; that which follows the application of organic matter lasts longer, more or less, according to its nature and the amount; but sooner or later the tentacles resume their former position, their glands glisten anew with fresh secretion, and they are ready to act again.

As to how the impulse is originated and propagated, and how the movements are made, comparatively simple as the structure is, we know as little as we do of the nature of nervous impulse and muscular motion. But two things Mr. Darwin has wellnigh made out, both of them by means and observations so simple and direct as to command our confidence, although they are contrary to the prevalent teaching. First, the transmission is through the ordinary cellular tissue, and not through what are called the fibrous or vascular bundles. Second, the movement is a vital one, and is effected by contraction on the side toward which the bending takes place, rather than by turgescent tension of the opposite side. The tentacle is pulled over rather than pushed over. So far all accords with muscular action.

The operation of this fly-catching apparatus, in any case, is plain. If the insect alights upon the disk of the leaf, the viscid secretion holds it fast—at least, an ordinary fly is unable to escape—its struggles only increase the number of glands involved and the amount of excitement; this is telegraphed to the surrounding and successively longer tentacles, which bent over in succession, so that within ten to thirty hours, if the leaf is active and the fly large enough, every one of the glands (on the average, nearly two hundred in number) will be found applied to the body of the insect. If the insect is small, and the lodgment toward one side, only the neighboring tentacles may take part in the capture. If two or three of the strong marginal tentacles are first encountered, their prompt inflection carries the intruder to the centre, and presses it down upon the glands which thickly pave the floor; these notify all the surrounding tentacles of the capture, that they may share the spoil, and the fate of that victim is even as of the first. A bit of meat or a crushed insect is treated in the same way.

This language implies that the animal matter is in some way or other discerned by the tentacles, and is appropriated. Formerly there was only a presumption of this, on the general ground that such an organization could hardly be purposeless. Yet, while such expressions were natural, if not unavoidable, they generally were used by those familiar with the facts in a half-serious, half-metaphorical sense. Thanks to Mr. Darwin's investigations, they may now be used in simplicity and seriousness.

That the glands secrete the glairy liquid of the drop is evident, not only from its nature, but from its persistence through a whole day's exposure to a summer sun, as also from its renewal after it has been removed, dried up, or absorbed. That they absorb as well as secrete, and that the whole tentacle may be profoundly affected thereby, are proved by the different effects, in kind and degree, which follow the application of different substances. Drops of rain-water, like single momentary touches of a solid body, produce no effect, as indeed they could be of no advantage; but a little carbonate of ammonia in the water, or an infusion of meat, not only causes inflection, but promptly manifests its action upon the contents of the cells of which the tentacle is constructed. These cells are sufficiently transparent to be viewed under the microscope without dissection or other interference; and the change which takes place in the fluid contents of these cells, when the gland above has been acted upon, is often visible through a weak lens, or sometimes even by the naked eye, although higher powers are required to discern what actually takes place. This change, which Mr. Darwin discovered, and turns to much account in his researches, he terms "aggregation of the protoplasm." When untouched and quiescent, the contents appear as an homogeneous purple fluid. When the gland is acted upon, minute purple particles appear, suspended in the now colorless or almost colorless fluid; and this change appears first in the cells next the gland, and then in those next beneath, traveling down the whole length of the tentacle. When the action is slight, this appearance does not last long; the particles of "aggregated protoplasm redissolved, the process of redissolution traveling upward from the base of the tentacle to the gland in a reverse direction to that of the aggregation. Whenever the action is more prolonged or intense, as when a bit of meat or crushed fly, or a fitting solution, is left upon the gland, the aggregation proceeds further, so that the whole protoplasm of each cell condenses into one or two masses, or into a single mass which will often separate into two, which afterward reunite; indeed, they incessantly change their forms and positions, being never at rest, although their movements are rather slow. In appearance and movements they are very like amoebae and the white corpuscles of the blood. Their motion, along with the streaming movement of rotation in the layer of white granular protoplasm that flows along the walls of the cell, under the high powers of the microscope "presents a wonderful scene of vital activity." This continues while the tentacle is inflected or the gland fed by animal matter, but vanishes by dissolution when the work is over and the tentacle straightens. That absorption takes place, and matter is conveyed from cell to cell, is well made out, especially by the experiments with carbonate of ammonia. Nevertheless, this aggregation is not dependent upon absorption, for it equally occurs from mechanical irritation of the gland, and always accompanies inflection, however caused, though it may take place without it. This is also apparent from the astonishingly minute quantity of certain substances which suffices to produce sensible inflection and aggregation—such, for instance, as the 1/20000000 or even the 1/30000000 of a grain of phosphate or nitrate of ammonia!

By varied experiments it was found that the nitrate of ammonia was more powerful than the carbonate, and the phosphate more powerful than the nitrate, this result being intelligible from the difference in the amount of nitrogen in the first two salts, and from the presence of phosphorus in the third. There is nothing surprising in the absorption of such extremely dilute solutions by a gland. As our author remarks: "All physiologists admit that the roots of plants absorb the salts of ammonia brought to them by the rain; and fourteen gallons of rain-water contain a grain of ammonia; therefore, only a little more than twice as much as in the weakest solution employed by me. The fact which appears truly wonderful is that the 1/20000000 of a grain of the phosphate of ammonia, including less than 1/30000000 of efficient matter (if the water of crystallization is deducted), when absorbed by a gland, should induce some change in it which leads to a motor impulse being transmitted down the whole length of the tentacle, causing its basal part to bend, often through an angle of 180 degrees." But odoriferous particles which act upon the nerves of animals must be infinitely smaller, and by these a dog a quarter of a mile to the leeward of a deer perceives his presence by some change in the olfactory nerves transmitted through them to the brain.

When Mr. Darwin obtained these results, fourteen years ago, he could claim for Drosera a power and delicacy in the detection of minute quantities of a substance far beyond the resources of the most skillful chemist; but in a foot-note he admits that "now the spectroscope has altogether beaten Drosera; for, according to Bunsen and Kirchhoff, probably less than the 1/200000000 of a grain of sodium can be thus detected."

Finally, that this highly-sensitive and active living organism absorbs, will not be doubted when it is proved to digest, that is, to dissolve otherwise insoluble animal matter by the aid of special secretions. That it does this is now past doubting. In the first place, when the glands are excited they pour forth an increased amount of the ropy secretion. This occurs directly when a bit of meat is laid upon the central glands; and the influence which they transmit to the long-stalked marginal glands causes them, while incurving their tentacles, to secrete more copiously long before they have themselves touched anything. The primary fluid, secreted without excitation, does not of itself digest. But the secretion under excitement changes in Nature and becomes acid. So, according to Schiff, mechanical irritation excites the glands of the stomach to secrete an acid. In both this acid appears to be necessary to, but of itself insufficient for, digestion. The requisite solvent, a kind of ferment called pepsin, which acts only in the presence of the acid, is poured forth by the glands of the stomach only after they have absorbed certain soluble nutritive substances of the food; then this pepsin promptly dissolves muscle, fibrine, coagulated albumen, cartilage, and the like. Similarly it appears that Drosera-glands, after irritation by particles of glass, did not act upon little cubes of albumen. But when moistened with saliva, or replaced by bits of roast-meat or gelatine, or even cartilage, which supply some soluble peptone-matter to initiate the process, these substances are promptly acted upon, and dissolved or digested; whence it is inferred that the analogy with the stomach holds good throughout, and that a ferment similar to pepsin is poured out under the stimulus of some soluble animal matter. But the direct evidence of this is furnished only by the related carnivorous plant, Dionaea, from which the secretions, poured out when digestion is about to begin, may be collected in quantity sufficient for chemical examination. In short, the experiments show "that there is a remarkable accordance in the power of digestion between the gastric juice of animals, with its pepsin and hydrochloric acid, and the secretion of Drosera, with its ferment and acid belonging to the acetic series. We can, therefore, hardly doubt that the ferment in both cases is closely similar, if not identically the same. That a plant and an animal should pour forth the same, or nearly the same, complex secretion, adapted for the same purpose of digestion, is a new and wonderful fact in physiology."

There are one or two other species of sundew—one of them almost as common in Europe and North America as the ordinary round-leaved species—which act in the same way, except that, having their leaves longer in proportion to their breadth, their sides never curl inward, but they are much disposed to aid the action of their tentacles by incurving the tip of the leaf, as if to grasp the morsel. There are many others, with variously less efficient and less advantageously arranged insectivorous apparatus, which, in the language of the new science, may be either on the way to acquire something better, or of losing what they may have had, while now adapting themselves to a proper vegetable life. There is one member of the family (Drosophyllum Lusitanicum), an almost shrubby plant, which grows on dry and sunny hills in Portugal and Morocco—which the villagers call "the flycatcher," and hang up in their cottages for the purpose—the glandular tentacles of which have wholly lost their powers of movement, if they ever had any, but which still secrete, digest, and absorb, being roused to great activity by the contact of any animal matter. A friend of ours once remarked that it was fearful to contemplate the amount of soul that could be called forth in a dog by the sight of a piece of meat. Equally wonderful is the avidity for animal food manifested by these vegetable tentacles, that can "only stand and wait" for it.

Only a brief chapter is devoted to Dionaea of North Carolina, the Venus's fly-trap, albeit, "from the rapidity and force of its movements, one of the most wonderful in the world." It is of the same family as the sundew; but the action is transferred from tentacles on the leaf to the body of the leaf itself, which is transformed into a spring-trap, closing with a sudden movement over the alighted insect. No secretion is provided beforehand either for allurement or detention; but after the captive is secured, microscopic glands within the surface of the leaf pour out an abundant gastric juice to digest it. Mrs. Glass's classical directions in the cook-book, "first catch your hare," are implicitly followed.

Avoiding here all repetition or recapitulation of our former narrative, suffice it now to mention two interesting recent additions to our knowledge, for which we are indebted to Mr. Darwin. One is a research, the other an inspiration. It is mainly his investigations which have shown that the glairy liquid, which is poured upon and macerates the captured insect, accomplishes a true digestion; that, like the gastric juice of animals, it contains both a free acid and pepsin or its analogue, these two together dissolving albumen, meat, and the like. The other point relates to the significance of a peculiarity in the process of capture. When the trap suddenly incloses an insect which has betrayed its presence by touching one of the internal sensitive bristles, the closure is at first incomplete. For the sides approach in an arching way, surrounding a considerable cavity, and the marginal spine-like bristles merely intercross their tips, leaving intervening spaces through which one may look into the cavity beneath. A good idea may be had of it by bringing the two palms near together to represent the sides of the trap, and loosely interlocking the fingers to represent the marginal bristles or bars. After remaining some time in this position the closure is made complete by the margins coming into full contact, and the sides finally flattening down so as to press firmly upon the insect within; the secretion excited by contact is now poured out, and digestion begins. Why these two stages? Why should time be lost by this preliminary and incomplete closing? The query probably was never distinctly raised before, no one noticing anything here that needed explanation. Darwinian teleology, however, raises questions like this, and Mr. Darwin not only propounded the riddle but solved it. The object of the partial closing is to permit small insects to escape through the meshes, detaining only those plump enough to be worth the trouble of digesting. For naturally only one insect is caught at a time, and digestion is a slow business with Dionaeas, as with anacondas, requiring ordinarily a fortnight. It is not worth while to undertake it with a gnat when larger game may be had. To test this happy conjecture, Mr. Canby was asked, on visiting the Dionaeas in their native habitat, to collect early in the season a good series of leaves in the act of digesting naturally-caught insects. Upon opening them it was found that ten out of fourteen were engaged upon relatively large prey, and of the remaining four three had insects as large as ants, and one a rather small fly.

"There be land-rats and water-rats" in this carnivorous sun-dew family. Aldrovanda, of the warmer parts of Europe and of India, is an aquatic plant, with bladdery leaves, which were supposed to be useful in rendering the herbage buoyant in water. But it has recently been found that the bladder is composed of two lobes, like the trap of its relative Dionaea, or the valves of a mussel-shell; that these open when the plant is in an active state, are provided with some sensitive bristles within, and when these are touched close with a quick movement. These water-traps are manifestly adapted for catching living creatures; and the few incomplete investigations that have already been made render it highly probably that they appropriate their prey for nourishment; whether by digestion or by mere absorption of decomposing animal matter, is uncertain. It is certainly most remarkable that this family of plants, wherever met with, and under the most diverse conditions and modes of life, should always in some way or other be predaceous and carnivorous.

If it be not only surprising but somewhat confounding to our classifications that a whole group of plants should subsist partly by digesting animal matter and partly in the normal way of decomposing carbonic acid and producing the basis of animal matter, we have, as Mr. Darwin remarks, a counterpart anomaly in the animal kingdom. While some plants have stomachs, some animals have roots. "The rhizocephalous crustaceans do not feed like other animals by their mouths, for they are destitute of an alimentary canal, but they live by absorbing through root-like processes the juices of the animals on which they are parasitic."

To a naturalist of our day, imbued with those ideas of the solidarity of organic Nature which such facts as those we have been considering suggest, the greatest anomaly of all would be that they are really anomalous or unique. Reasonably supposing, therefore, that the sundew did not stand alone, Mr. Darwin turned his attention to other groups of plants; and, first, to the bladderworts, which have no near kinship with the sundews, but, like the aquatic representative of that family, are provided with bladdery sacs, under water. In the common species of Utricularia or bladderwort, these little sacs, hanging from submerged leaves or branches, have their orifice closed by a lid which opens inwardly—a veritable trapdoor. It had been noticed in England and France that they contained minute crustacean animals. Early in the summer of 1874, Mr. Darwin ascertained the mechanism for their capture and the great success with which it is used. But before his account was written out, Prof. Cohn published an excellent paper on the subject in Germany; and Mrs. Treat, of Vineland, New Jersey, a still earlier one in this country—in the New York Tribune in the autumn of 1874. Of the latter, Mr. Darwin remarks that she "has been more successful than any other observer in witnessing the actual entrance of these minute creatures." They never come out, but soon perish in their prison, which receives a continued succession of victims, but little, if any, fresh air to the contained water. The action of the trap is purely mechanical, without evident irritability in the opening or shutting. There is no evidence nor much likelihood of proper digestion; indeed, Mr. Darwin found evidence to the contrary. But the more or less decomposed and dissolved animal matter is doubtless absorbed into the plant; for the whole interior of the sac is lined with peculiar, elongated and four-armed very thin-walled processes, which contain active protoplasm, and which were proved by experiment to "have the power of absorbing matter from weak solutions of certain salts of ammonia and urea, and from a putrid infusion of raw meat."

Although the bladderworts "prey on garbage," their terrestrial relatives "live cleanly," as nobler plants should do, and have a good and true digestion. Pinguicula, or butterwort, is the representative of this family upon land. It gets both its Latin and its English name from the fatty or greasy appearance of the upper face of its broad leaves; and this appearance is due to a dense coat or pile of short-stalked glands, which secrete a colorless and extremely viscid liquid. By this small flies, or whatever may alight or fall upon the leaf, are held fast. These waifs might be useless or even injurious to the plant. Probably Mr. Darwin was the first to ask whether they might be of advantage. He certainly was the first to show that they probably are so. The evidence from experiment, shortly summed up, is, that insects alive or dead, and also other nitrogenous bodies, excite these glands to increased secretion; the secretion then becomes acid, and acquires the power of dissolving solid animal substances—that is, the power of digestion in the manner of Drosera and Dionaea. And the stalks of their glands under the microscope give the same ocular evidence of absorption. The leaves of the butterwort are apt to have their margins folded inward, like a rim or hem. Taking young and vigorous leaves to which hardly anything had yet adhered, and of which the margins were still flat, Mr. Darwin set within one margin a row of small flies. Fifteen hours afterward this edge was neatly turned inward, partly covering the row of flies, and the surrounding glands were secreting copiously. The other edge remained flat and unaltered. Then he stuck a fly to the middle of the leaf just below its tip, and soon both margins infolded, so as to clasp the object. Many other and varied experiments yielded similar results. Even pollen, which would not rarely be lodged upon these leaves, as it falls from surrounding wind-fertilized plants, also small seeds, excited the same action, and showed signs of being acted upon. "We may therefore conclude," with Mr. Darwin, "that Pinguicula vulgaris, with its small roots, is not only supported to a large extent by the extraordinary number of insects which it habitually captures, but likewise draws some nourishment from the pollen, leaves, and seeds, of other plants which often adhere to its leaves. It is, therefore, partly a vegetable as well as an animal feeder."

What is now to be thought of the ordinary glandular hairs which render the surface of many and the most various plants extremely viscid? Their number is legion. The Chinese primrose of common garden and house culture is no extraordinary instance; but Mr. Francis Darwin, counting those on a small space measured by the micrometer, estimated them at 65,371 to the square inch of foliage, taking in both surfaces of the leaf, or two or three millions on a moderate-sized specimen of this small herb. Glands of this sort were loosely regarded as organs for excretion, without much consideration of the question whether, in vegetable life, there could be any need to excrete, or any advantage gained by throwing off such products; and, while the popular name of catch-fly, given to several common species of Silene, indicates long familiarity with the fact, probably no one ever imagined that the swarms of small insects which perish upon these sticky surfaces were ever turned to account by the plant. In many such cases, no doubt they perish as uselessly as when attracted into the flame of a candle. In the tobacco-plant, for instance, Mr. Darwin could find no evidence that the glandular hairs absorb animal matter. But Darwinian philosophy expects all gradations between casualty and complete adaptation. It is most probable that any thin-walled vegetable structure which secretes may also be capable of absorbing under favorable conditions. The myriads of exquisitely-constructed glands of the Chinese primrose are not likely to be functionless. Mr. Darwin ascertained by direct experiment that they promptly absorb carbonate of ammonia, both in watery solution and in vapor. So, since rain-water usually contains a small percentage of ammonia, a use for these glands becomes apparent—one completely congruous with that of absorbing any animal matter, or products of its decomposition, which may come in their way through the occasional entanglement of insects in their viscid secretion. In several saxifrages—not very distant relatives of Drosera—the viscid glands equally manifested the power of absorption.

To trace a gradation between a simply absorbing hair with a glutinous tip, through which the plant may perchance derive slight contingent advantage, and the tentacles of a sundew, with their exquisite and associated adaptations, does not much lessen the wonder nor explain the phenomena. After all, as Mr. Darwin modestly concludes, "we see how little has been made out in comparison with what remains unexplained and unknown." But all this must be allowed to be an important contribution to the doctrine of the gradual acquirement of uses and functions, and hardly to find conceivable explanation upon any other hypothesis.

There remains one more mode in which plants of the higher grade are known to prey upon animals; namely, by means of pitchers, urns, or tubes, in which insects and the like are drowned or confined, and either macerated or digested. To this Mr. Darwin barely alludes on the last page of the present volume. The main facts known respecting the American pitcher-plants have, as was natural, been ascertained in this country; and we gave an abstract, two years ago, of our then incipient knowledge. Much has been learned since, although all the observations have been of a desultory character. If space permitted, an instructive narrative might be drawn up, as well of the economy of the Sarracenias as of how we came to know what we do of it. But the very little we have room for will be strictly supplementary to our former article.

The pitchers of our familiar Northern Sarracenia, which is likewise Southern, are open-mouthed; and, although they certainly secrete some liquid when young, must derive most of the water they ordinarily contain from rain. How insects are attracted is unknown, but the water abounds with their drowned bodies and decomposing remains.

In the more southern S. flava, the long and trumpet-shaped pitchers evidently depend upon the liquid which they themselves secrete, although at maturity, when the hood becomes erect, rain may somewhat add to it. This species, as we know, allures insects by a peculiar sweet exudation within the orifice; they fall in and perish, though seldom by drowning, yet few are able to escape; and their decomposing remains accumulate in the narrow bottom of the vessel. Two other long-tubed species of the Southern States are similar in these respects. There is another, S. psittacina, the parrot-headed species, remarkable for the cowl-shaped hood so completely inflexed over the mouth of the small pitcher that no rain can possibly enter. Little is known, however, of the efficiency of this species as a fly-catcher; but its conformation has a morphological interest, leading up, as it does, to the Californian type of pitcher presently to be mentioned.

But the remaining species, S. variolaris, is the most wonderful of our pitcher-plants in its adaptations for the capture of insects. The inflated and mottled lid or hood overarches the ample orifice of the tubular pitcher sufficiently to ward off the rain, but not to obstruct the free access of flying insects. Flies, ants, and most insects, glide and fall from the treacherous smooth throat into the deep well below, and never escape. They are allured by a sweet secretion just within the orifice— which was discovered and described long ago, and the knowledge of it wellnigh forgotten until recently. And, finally, Dr. Mellichamp, of South Carolina, two years ago made the capital discovery that, during the height of the season, this lure extends from the orifice down nearly to the ground, a length of a foot or two, in the form of a honeyed line or narrow trail on the edge of the wing-like border which is conspicuous in all these species, although only in this one, so far as known, turned to such account. Here, one would say, is a special adaptation to ants and such terrestrial and creeping insects. Well, long before this sweet trail was known, it was remarked by the late Prof. Wyman and others that the pitchers of this species, in the savannahs of Georgia and Florida, contain far more ants than they do of all other insects put together.

Finally, all this is essentially repeated in the peculiar Californian pitcher-plant (Darlingtonia), a genus of the same natural family, which captures insects in great variety, enticing them by a sweetish secretion over the whole inside of the inflated hood and that of a curious forked appendage, resembling a fish-tail, which overhangs the orifice. This orifice is so concealed that it can be seen and approached only from below, as if—the casual observer might infer—to escape visitation. But dead insects of all kinds, and their decomposing remains, crowd the cavity and saturate the liquid therein contained, enticed, it is said, by a peculiar odor, as well as by the sweet lure which is at some stages so abundant as to drip from the tips of the overhanging appendage. The principal observations upon this pitcher-plant in its native habitat have been made by Mrs. Austin, and only some of the earlier ones have thus far been published by Mr. Canby. But we are assured that in this, as in the Sarracenia variolaris, the sweet exudation extends at the proper season from the orifice down the wing nearly to the ground, and that ants follow this honeyed pathway to their destruction. Also, that the watery liquid in the pitcher, which must be wholly a secretion, is much increased in quantity after the capture of insects.

It cannot now well be doubted that the animal matter is utilized by the plant in all these cases, although most probably only after maceration or decomposition. In some of them even digestion, or at least the absorption of undecomposed soluble animal juices, may be suspected; but there is no proof of it. But, if pitchers of the Sarracenia family are only macerating vessels, those of Nepenthes—the pitchers of the Indian Archipelago, familiar in conservatories—seem to be stomachs. The investigations of the President of the Royal Society, Dr. Hooker, although incomplete, wellnigh demonstrate that these not only allure insects by a sweet secretion at the rim and upon the lid of the cup, but also that their capture, or the presence of other partly soluble animal matter, produces an increase and an acidulation of the contained watery liquid, which thereupon becomes capable of acting in the manner of that of Drosera and Dionaea, dissolving flesh, albumen, and the like.

After all, there never was just ground for denying to vegetables the use of animal food. The fungi are by far the most numerous family of plants, and they all live upon organic matter, some upon dead and decomposing, some upon living, some upon both; and the number of those that feed upon living animals is large. Whether these carnivorous propensities of higher plants which so excite our wonder be regarded as survivals of ancestral habits, or as comparatively late acquirements, or even as special endowments, in any case what we have now learned of them goes to strengthen the conclusion that the whole organic world is akin.

The volume upon "The Movements and Habits of Climbing Plants" is a revised and enlarged edition of a memoir communicated to the Linnaean Society in 1865, and published in the ninth volume of its Journal. There was an extra impression, but, beyond the circle of naturalists, it can hardly have been much known at first-hand. Even now, when it is made a part of the general Darwinian literature, it is unlikely to be as widely read as the companion volume which we have been reviewing; although it is really a more readable book, and well worthy of far more extended notice at our hands than it can now receive. The reason is obvious. It seems as natural that plants should climb as it does unnatural that any should take animal food. Most people, knowing that some plants "twine with the sun," and others "against the sun," have an idea that the sun in some way causes the twining; indeed, the notion is still fixed in the popular mind that the same species twines in opposite directions north and south of the equator.

Readers of this fascinating treatise will learn, first of all, that the sun has no influence over such movements directly, and that its indirect influence is commonly adverse or disturbing, except the heat, which quickens vegetable as it does animal life. Also, that climbing is accomplished by powers and actions as unlike those generally predicated of the vegetable kingdom as any which have been brought to view in the preceding volume. Climbing plants "feel" as well as "grow and live;" and they also manifest an automatism which is perhaps more wonderful than a response by visible movement to an external irritation. Nor do plants grow up their supports, as is unthinkingly supposed; for, although only growing or newly-grown parts act in climbing, the climbing and the growth are entirely distinct. To this there is one exception—an instructive one, as showing how one action passes into another, and how the same result may be brought about in different ways—that of stems which climb by rootlets, such as of ivy and trumpet-creeper. Here the stem ascends by growth alone, taking upward direction, and is fixed by root-lets as it grows. There is no better way of climbing walls, precipices, and large tree-trunks.

But small stems and similar supports are best ascended by twining; and this calls out powers of another and higher order. The twining stem does not grow around its support, but winds around it, and it does this by a movement the nature of which is best observed in stems which have not yet reached their support, or have overtopped it and stretched out beyond it. Then it may be seen that the extending summit, reaching farther and farther as it grows, is making free circular sweeps, by night as well as by day, and irrespective of external circumstances, except that warmth accelerates the movement, and that the general tendency of young stems to bend toward the light may, in case of lateral illumination, accelerate one-half the circuit while it equally retards the other. The arrest of the revolution where the supporting body is struck, while the portion beyond continues its movement, brings about the twining. As to the proximate cause of this sweeping motion, a few simple experiments prove that it results from the bowing or bending of the free summit of the stem into a more or less horizontal position (this bending being successively to every point of the compass, through an action which circulates around the stem in the direction of the sweep), and of the consequent twining, i.e., "with the sun," or with the movement of the hands of a watch, in the hop, or in the opposite direction in pole-beans and most twiners. Twining plants, therefore, ascend trees or other stems by an action and a movement of their own, from which they derive advantage. To plants liable to be overshadowed by more robust companions, climbing is an economical method of obtaining a freer exposure to light and air with the smallest possible expenditure of material. But twiners have one disadvantage: to rise ten feet they must produce fifteen feet of stem or thereabouts, according to the diameter of the support, and the openness or closeness of the coil. A rootlet-climber saves much in this respect, but has a restricted range of action, and other disadvantages.

There are two other modes, which combine the utmost economy of material with freer range of action. There are, in the first place, leaf-climbers of various sorts, agreeing only in this, that the duty of laying hold is transferred to the leaves, so that the stem may rise in a direct line. Sometimes the blade or leaflets, or some of them, but more commonly their slender stalks, undertake the work, and the plant rises as a boy ascends a tree, grasping first with one hand or arm, then with the other. Indeed, the comparison, like the leaf-stalk, holds better than would be supposed; for the grasping of the latter is not the result of a blind groping in all directions by a continuous movement, but of a definite sensitiveness which acts only upon the occasion. Most leaves make no regular sweeps; but when the stalks of a leaf-climbing species come into prolonged contact with any fitting extraneous body, they slowly incurve and make a turn around it, and then commonly thicken and harden until they attain a strength which may equal that of the stem itself. Here we have the faculty of movement to a definite end, upon external irritation, of the same nature with that displayed by Dionaea and Drosera, although slower for the most part than even in the latter. But the movement of the hour-hand of the clock is not different in nature or cause from that of the second-hand.

Finally—distribution of office being, on the whole, most advantageous and economical, and this, in the vegetable kingdom, being led up to by degrees—we reach, through numerous gradations, the highest style of climbing plants in the tendril-climber. A tendril morphologically, is either a leaf or branch of stem, or a portion of one, specially organized for climbing. Some tendrils simply turn away from light, as do those of grape-vines, thus taking the direction in which some supporting object is likely to be encountered; most are indifferent to light; and many revolve in the manner of the summit of twining stems. As the stems which bear these highly-endowed tendrils in many cases themselves also revolve more or less, though they seldom twine, their reach is the more extensive; and to this endowment of automatic movement most tendrils add the other faculty, that of incurving and coiling upon prolonged touch, or even brief contact, in the highest degree. Some long tendrils, when in their best condition, revolve so rapidly that the sweeping movement may be plainly seen; indeed, we have seen a quarter-circuit in a Passiflora sicyoides accomplished in less than a minute, and the half-circuit in ten minutes; but the other half (for a reason alluded to in the next paragraph) takes a much longer time. Then, as to the coiling upon contact, in the case first noticed in this country,[XI-3] in the year 1858, which Mr. Darwin mentions as having led him into this investigation, the tendril of Sicyos was seen to coil within half a minute after a stroke with the hand, and to make a full turn or more within the next minute; furnishing ocular evidence that tendrils grasp and coil in virtue of sensitiveness to contact, and, one would suppose, negativing Sachs's recent hypothesis that all these movements are owing "to rapid growth on the side opposite to that which becomes concave"—a view to which Mr. Darwin objects, but not so strongly as he might. The tendril of this sort, on striking some fitting object, quickly curls round and firmly grasps it; then, after some hours, one side shortening or remaining short in proportion to the other, it coils into a spire, dragging the stem up to its support, and enabling the next tendril above to secure a readier hold.

In revolving tendrils perhaps the most wonderful adaptation is that by which they avoid attachment to, or winding themselves upon, the ascending summit of the stem that bears them. This they would inevitably do if they continued their sweep horizontally. But when in its course it nears the parent

stem the tendril moves slowly, as if to gather strength, then C.~ stiffens and rises into an erect position parallel with it, and C so passes by the dangerous point; after which it comes rapidly down to the horizontal position, in which it moves until it again approaches and again avoids the impending obstacle.

Climbing plants are distributed throughout almost all the natural orders. In some orders climbing is the rule, in most it is the exception, occurring only in certain genera. The tendency of stems to move in circuits—upon which climbing more commonly depends, and out of which it is conceived to have been educed—is manifested incipiently by many a plant which does not climb. Of those that do there are all degrees, from the feeblest to the most efficient, from those which have no special adaptation to those which have exquisitely-endowed special organs for climbing. The conclusion reached is, that the power "is inherent, though undeveloped, in almost every plant;" "that climbing plants have utilized and perfected a widely-distributed and incipient capacity, which, as far as we can see, is of no service to ordinary plants."

Inherent powers and incipient manifestations, useless to their possessors but useful to their successors—this, doubtless, is according to the order of Nature; but it seems to need something more than natural selection to account for it.

XII

DURATION AND

ORIGINATION OF

RACE AND SPECIES—

IMPORT OF SEXUAL REPRODUCTION

I Do Varieties wear out, or tend to wear out?

(New York Tribune, and American Journal of Science and the Arts, February, 1875)

This question has been argued from time to time for more than half a century, and is far from being settled yet. Indeed, it is not to be settled either way so easily as is sometimes thought. The result of a prolonged and rather lively discussion of the topic about forty years ago in England, in which Lindley bore a leading part on the negative side, was, if we rightly remember, that the nays had the best of the argument. The deniers could fairly well explain away the facts adduced by the other side, and evade the force of the reasons then assigned to prove that varieties were bound to die out in the course of time. But if the case were fully re-argued now, it is by no means certain that the nays would win it. The most they could expect would be the Scotch verdict, "not proven." And this not because much, if any, additional evidence of the actual wearing out of any variety has turned up since, but because a presumption has been raised under which the evidence would take a bias the other way. There is now in the minds of scientific men some reason to expect that certain varieties would die out in the long run, and this might have an important influence upon the interpretation of the facts. Curiously enough, however, the recent discussions to which our attention has been called seem, on both sides, to have overlooked this.

But, first of all, the question needs to be more specifically stated. There are varieties and varieties. They may, some of them, disappear or deteriorate, but yet not wear out—not come to an end from any inherent cause. One might even say, the younger they are the less the chance of survival unless well cared for. They may be smothered out by the adverse force of superior numbers; they are even more likely to be bred out of existence by unprevented cross-fertilization, or to disappear from mere change of fashion. The question, however, is not so much about reversion to an ancestral state, or the falling off of a high-bred stock into an inferior condition. Of such cases it is enough to say that, when a variety or strain, of animal or vegetable, is led up to unusual fecundity or of size or product of any organ, for our good, and not for the good of the plant or animal itself, it can be kept so only by high feeding and exceptional care; and that with high feeding and artificial appliances comes vastly increased liability to disease, which may practically annihilate the race. But then the race, like the bursted boiler, could not be said to wear out, while if left to ordinary conditions, and allowed to degenerate back into a more natural if less useful state, its hold on life would evidently be increased rather than diminished.

As to natural varieties or races under normal conditions, sexually propagated, it could readily be shown that they are neither more nor less likely to disappear from any inherent cause than the species from which they originated. Whether species wear out, i.e., have their rise, culmination, and decline, from any inherent cause, is wholly a geological and very speculative problem, upon which, indeed, only vague conjectures can be offered. The matter actually under discussion concerns cultivated domesticated varieties only, and, as to plants, is covered by two questions.

First, Will races propagated by seed, being so fixed that they come true to seed, and purely bred (not crossed with any other sort), continue so indefinitely, or will they run out in time—not die out, perhaps, but lose their distinguishing characters? Upon this, all we are able to say is that we know no reason why they should wear out or deteriorate from any inherent cause. The transient existence or the deterioration and disappearance of many such races are sufficiently accounted for otherwise; as in the case of extraordinarily exuberant varieties, such as mammoth fruits or roots, by increased liability to disease, already adverted to, or by the failure of the high feeding they demand. A common cause, in ordinary cases, is cross-breeding, through the agency of wind or insects, which is difficult to guard against. Or they go out of fashion and are superseded by others thought to be better, and so the old ones disappear.

Or, finally, they may revert to an ancestral form. As offspring tend to resemble grandparents almost as much as parents, and as a line of close-bred ancestry is generally prepotent, so newly-originated varieties have always a tendency to reversion. This is pretty sure to show itself in some of the progeny of the earlier generations, and the breeder has to guard against it by rigid selection. But the older the variety is—that is, the longer the series of generations in which it has come true from seed—the less the chance of reversion: for now, to be like the immediate parents, is also to be like a long line of ancestry; and so all the influences concerned—- that is, both parental and ancestral heritability—act in one and the same direction. So, since the older a race is the more reason it has to continue true, the presumption of the unlimited permanence of old races is very strong.

Of course the race itself may give off new varieties; but that is no interference with the vitality of the original stock. If some of the new varieties supplant the old, that will not be because the unvaried stock is worn out or decrepit with age, but because in wild Nature the newer forms are better adapted to the surroundings, or, under man's care, better adapted to his wants or fancies.

The second question, and one upon which the discussion about the wearing out of varieties generally turns, is, Will varieties propagated from buds, i.e., by division, grafts, bulbs, tubers, and the like, necessarily deteriorate and die out? First, Do they die out as a matter of fact? Upon this, the testimony has all along been conflicting. Andrew Knight was sure that they do, and there could hardly be a more trustworthy witness.

"The fact," he says, fifty years ago, "that certain varieties of some species of fruit which have been long cultivated cannot now be made to grow in the same soils and under the same mode of management, which was a century ago so perfectly successful, is placed beyond the reach of controversy. Every experiment which seemed to afford the slightest prospect of success was tried by myself and others to propagate the old varieties of the apple and pear which formerly constituted the orchards of Herefordshire, without a single healthy or efficient tree having been obtained; and I believe all attempts to propagate these varieties have, during some years, wholly ceased to be made."

To this it was replied, in that and the next generation, that cultivated vines have been transmitted by perpetual division from the time of the Romans, and that several of the sorts, still prized and prolific, are well identified, among them the ancient Graecula, considered to be the modern Corinth or currant grape, which has immemorially been seedless; that the old nonpareil apple was known in the time of Queen Elizabeth; that the white beurre pears of France have been propagated from the earliest times; and that golden pippins, St. Michael pears, and others said to have run out, were still to be had in good condition.

Coming down to the present year, a glance through the proceedings of pomological societies, and the debates of farmers' clubs, brings out the same difference of opinion. The testimony is nearly equally divided. Perhaps the larger number speak of the deterioration and failure of particular old sorts; but when the question turns on "wearing out," the positive evidence of vigorous trees and sound fruits is most telling. A little positive testimony outweighs a good deal of negative. This cannot readily be explained away, while the failures may be, by exhaustion of soil, incoming of disease, or alteration of climate or circumstances. On the other hand, it may be urged that, if a variety of this sort is fated to become decrepit and die out, it is not bound to die out all at once, and everywhere at the same time. It would be expected first to give way wherever it is weakest, from whatever cause. This consideration has an important bearing upon the final question, Are old varieties of this kind on the way to die out on account of their age or any inherent limit of vitality?

Here, again, Mr. Knight took an extreme view. In his essay in the "Philosophical Transactions," published in the year 1810, he propounded the theory, not merely of a natural limit to varieties from grafts and cuttings, but even that they would not survive the natural term of the life of the seedling trees from which they were originally taken. Whatever may have been his view of the natural term of the life of a tree, and of a cutting being merely a part of the individual that produced it, there is no doubt that he laid himself open to the effective replies which were made from all sides at the time, and have lost none of their force since. Weeping-willows, bread-fruits, bananas, sugar-cane, tiger-lilies, Jerusalem artichokes, and the like, have been propagated for a long while in this way, without evident decadence. Moreover, the analogy upon which his hypothesis is founded will not hold. Whether or not one adopts the present writer's conception, that individuality is not actually reached or maintained in the vegetable world, it is clear enough that a common plant or tree is not an individual in the sense that a horse or man, or any one of the higher animals, is—that it is an individual only in the sense that a branching zoophyte or mass of coral is. Solvitur crescendo: the tree and the branch equally demonstrate that they are not individuals, by being divided with impunity and advantage, with no loss of life, but much increase. It looks odd enough to see a writer like Mr. Sisley reproducing the old hypothesis in so bare a form as this: "I am prepared to maintain that varieties are individuals, and that as they are born they must die, like other individuals . . . We know that oaks, Sequoias, and other trees, live several centuries, but how many we do not exactly know. But that they must die, no one in his senses will dispute." Now, what people in their senses do dispute is, not that the tree will die, but that other trees, established from its cuttings, will die with it.

But does it follow from this that non-sexually-propagated varieties are endowed with the same power of unlimited duration that is possessed by varieties and species propagated sexually—i.e., by seed? Those who think so jump too soon at their conclusion. For, as to the facts, it is not enough to point out the diseases or the trouble in the soil or the atmosphere to which certain old fruits are succumbing, nor to prove that a parasitic fungus (Peronospora infestans) is what is the matter with potatoes. For how else would constitutional debility, if such there be, more naturally manifest itself than in such increased liability or diminished resistance to such attacks? And if you say that, anyhow, such varieties do not die of old age—meaning that each individual attacked does not die of old age, but of manifest disease—it may be asked in return, what individual man ever dies of old age in any other sense than of a similar inability to resist invasions which in earlier years would have produced no noticeable effect? Aged people die of a slight cold or a slight accident, but the inevitable weakness that attends old age is what makes these slight attacks fatal.

Finally, there is a philosophical argument which tells strongly for some limitation of the duration of non-sexually propagated forms, one that probably Knight never thought of, but which we should not have expected recent writers to overlook. When Mr. Darwin announced the principle that cross-fertilization between the individuals of a species is the plan of Nature, and is practically so universal that it fairly sustains his inference that no hermaphrodite species continually self-fertilized would continue to exist, he made it clear to all who apprehend and receive the principle that a series of plants propagated by buds only must have weaker hold of life than a series reproduced by seed. For the former is the closest possible kind of close breeding. Upon this ground such varieties may be expected ultimately to die out; but "the mills of the gods grind so exceeding slow" that we cannot say that any particular grist has been actually ground out under human observation.

If it be asked how the asserted principle is proved or made probable, we can here merely say that the proof is wholly inferential. But the inference is drawn from such a vast array of facts that it is wellnigh irresistible. It is the legitimate explanation of those arrangements in Nature to secure cross-fertilization in the species, either constantly or occasionally, which are so general, so varied and diverse, and, we may add, so exquisite and wonderful, that, once propounded, we see that it must be true.* What else, indeed, is the meaning and

* Here an article would be in place, explaining the arrangements in Nature for cross-fertilization, or wide-breeding, in plants, through the agency, sometimes of the winds, but more commonly of insects; the more so, since the development of the principle, the appreciation of its importance, and its confirmation by abundant facts, are mainly due to Mr. Darwin. But our reviews and notices of his early work "On the Contrivances in Nature for the Fertilization of Orchids by Means of Insects, in 1862, and his various subsequent papers upon other parts of this subject, are either too technical or too fragmentary or special to be here reproduced. Indeed, a popular essay is now hardly needed, since the topic has been fully presented, of late years, in the current popular and scientific journals, and in common educational works and text-books, so that it is in the way of becoming a part—and a most inviting part—of ordinary botanical instruction. use of sexual reproduction? Not simply increase of numbers; for that is otherwise effectually provided for by budding propagation in plants and many of the lower animals. There are plants, indeed, of the lower sort (such as diatoms), in which the whole multiplication takes place in this way, and with great rapidity. These also have sexual reproduction; but in it two old individuals are always destroyed to make a single new one! Here propagation diminishes the number of individuals fifty per cent. Who can suppose that such a costly process as this, and that all the exquisite arrangements for cross-fertilization in hermaphrodite plants, do not subserve some most important purpose? How and why the union of two organisms, or generally of two very minute portions of them, should reenforce vitality, we do not know, and can hardly conjecture. But this must be the meaning of sexual reproduction.

The conclusion of the matter, from the scientific point of view, is, that sexually-propagated varieties or races, although liable to disappear through change, need not be expected to wear out, and there is no proof that they do; but, that non-sexually propagated varieties, though not especially liable to change, may theoretically be expected to wear out, but to be a very long time about it.

II

Do Species wear out? and if not, why not?

The question we have just been considering was merely whether races are, or may be, as enduring as species. As to the inherently unlimited existence of species themselves, or the contrary, this, as we have said, is a geological and very speculative problem. Not a few geologists and naturalists, however, have concluded, or taken for granted, that species have a natural term of existence—that they culminate, decline, and disappear through exhaustion of specific vitality, or some equivalent internal cause. As might be expected from the nature of the inquiry, the facts which bear upon the question are far from decisive. If the fact that species in general have not been interminable, but that one after another in long succession has become extinct, would seem to warrant this conclusion, the persistence through immense periods of no inconsiderable number of the lower forms of vegetable and animal life, and of a few of the higher plants from the Tertiary period to the present, tells even more directly for the limitless existence of species. The disappearance is quite compatible with the latter view; while the persistence of any species is hardly explicable upon any other. So that, even under the common belief of the entire stability and essential inflexibility of species, extinction is more likely to have been accidental than predetermined, and the doctrine of inherent limitation is unsupported by positive evidence.

On the other hand, it is an implication of the Darwinian doctrine that species are essentially unlimited in existence. When they die out—as sooner or later any species may—the verdict must be accidental death, under stress of adverse circumstances, not exhaustion of vitality; and, commonly, when the species seems to die out, it will rather have suffered change. For the stock of vitality which enables it to vary and. survive in changed forms under changed circumstances must be deemed sufficient for a continued unchanged existence under unaltered conditions. And, indeed, the advancement from simpler to more complex, which upon the theory must have attended the diversification, would warrant or require the supposition of increase instead of diminution of power from age to age.

The only case we call to mind which, under the Darwinian view, might be interpreted as a dying out from inherent causes, is that of a species which refuses to vary, and thus lacks the capacity of adaptation to altering conditions. Under altering conditions, this lack would be fatal. But this would be the fatality of some species or form in particular, not of species or forms generally, which, for the most part, may and do vary sufficiently, and in varying survive, seemingly none the worse, but rather the better, for their long tenure of life.

The opposite idea, however, is maintained by M. Naudin,[XII-1] in a detailed exposition of his own views of evolution, which differ widely from those of Darwin in most respects, and notably in excluding that which, in our day, gives to the subject its first claim to scientific (as distinguished from purely speculative) attention; namely, natural selection. Instead of the causes or operations collectively personified under this term, and which are capable of exact or probable appreciation, M. Naudin invokes "the two principles of rhythm and of the decrease of forces in Nature." He is a thorough evolutionist, starting from essentially the same point with Darwin; for he conceives of all the forms or species of animals and plants "comme tire tout entier d'un protoplasma primordial, uniform, instable, eminemment plastique." Also in "l'integration croissante de la force evolutive a mesure qu'elle se partage dans les formes produites, et la decroissance proportionelle de la plasticite de ces formes a mesure qu'elles s'eloignent davantage de leur origine, et qu'elles sont mieux arretees." As they get older, they gain in fixity through the operation of the fundamental law of inheritance; but the species, like the individual, loses plasticity and vital force. To continue in the language of the original:

"C'est dire qu'il y a eu, pour l'ensemble du monde organique, une periode de formation ou tout etait changeant et mobile, une phase analogue a la vie embryonnaire et a la jeunesse de chaque etre particulier; et qu'a cet age de mobilite et de croissance a succede une periode de stabilite, au moins relative, une sorte d'age adulte, ou la force evolutive, ayant acheve son oeuvre, n'est plus occupee qu'a la maintenir, sans pouvoir produire d'organismes nouveaux. Limitee en quantite, comme toutes les forces en jeu dans une planete ou dans un systeme sideral tout entier, cette force n'a pu accomplir qu'un travail limite; et du meme qu'un organisme, animal ou vegetal, ne croit pas indefiniment et qu'il s'arrete a des proportions que rien ne peut faire depasser, de meme aussi l'organisme total de la nature s'est arrete a un etat d'equilibre, dont la duree, selon toutes vraisemblances, doit etre beaucoup plus longue que celle de la phase de developpement et de croissance.

A fixed amount of "evolutive force" is given, to begin with. At first enormous, because none has been used up in work, it is necessarily enfeebled in the currents into which the stream divides, and the narrower and narrower channels in which it flows with slowly-diminishing power. Hence the limited although very unequal duration of all individuals, of all species, and of all types of organization. A multitude of forms have disappeared already, and the number of species, far from increasing, as some have believed, must, on the contrary, be diminishing. Some species, no doubt, have suffered death by violence or accident, by geological changes, local alteration of the conditions, or the direct or indirect attacks of other species; but these have only anticipated their fate, for M. Naudin contends that most of the extinct species have died a natural death from exhaustion of force, and that all the survivors are on the way to it. The great timepiece of Nature was wound up at the beginning, and is running down. In the earlier stages of great plasticity and exuberant power, diversification took place freely, but only in definite lines, and species and types multiplied. As the power of survival is inherently limited, still more the power of change: this diminishes in time, if we rightly apprehend the idea, partly through the waning of vital force, partly through the fixity acquired by heredity—like producing like, the more certainly in proportion to the length and continuity of the ancestral chain And so the small variations of species which we behold are the feeble remnants of the pristine plasticity and an exhausted force.[XII-2] This force of variation or origination of forms has acted rhythmically or intermittently, because each movement was the result of the rupture of an equilibrium, the liberation of a force which till then was retained in a potential state by some opposing force or obstacle, overcoming which it passes to a new equilibrium and so on Hence alternations of dynamic activity and static repose, of origination of species and types, alternated with periods of stability or fixity. The timepiece does not run down regularly, but "la force procede par saccades; et . . . par pulsations d'autant plus energiques que la nature etait plus pres de son commencement."

Such is the hypothesis. For a theory of evolution, this is singularly unlike Darwin's in most respects, and particularly in the kind of causes invoked and speculations indulged in. But we are not here to comment upon it beyond the particular point under consideration, namely, its doctrine of the inherently limited duration of species. This comes, it will be noticed, as a deduction from the modern physical doctrine of the equivalence of force. The reasoning is ingenious, but, if we mistake not, fallacious.

To call that "evolutive force" which produces the change of one kind of plant or animal into another, is simple and easy, but of little help by way of explanation. To homologize it with physical force, as M. Naudin's argument requires, is indeed a step, and a hardy one; but it quite invalidates the argument. For, if the "evolutive force" is a part of the physical force of the universe, of which, as he reminds us, the sum is fixed and the tendency is toward a stable equilibrium in which all change is to end, then this evolutive was derived from the physical force; and why not still derivable from it? What is to prevent its replenishment in vegetation, pari passu with that great operation in which physical force is stored up in vegetable organisms, and by the expenditure or transformation of which their work, and that of all animals, is carried on? Whatever be the cause (if any there be) which determines the decadence and death of species, one cannot well believe that it is a consequence of a diminution of their proper force by plant-development and division; for instance, that the sum of what is called vital force in a full-grown tree is not greater, instead of less, than that in the seeding, and in the grove greater than in the single parental tree. This power, if it be properly a force, is doubtless as truly derived from the sunbeam as is the power which the plant and animal expend in work. Here, then, is a source of replenishment as lasting as the sun itself, and a ground—so far as a supply of force is concerned—for indefinite duration. For all that any one can mean by the indefinite existence of species is, that they may (for all that yet appears) continue while the external conditions of their being or well-being continue.

Perhaps, however, M. Naudin does not mean that "evolutive force," or the force of vitality, is really homologous with common physical force, but only something which may be likened to it. In that case the parallel has only a metaphorical value, and the reason why variation must cease and species die out is still to seek. In short, if that which continues the series of individuals in propagation, whether like or unlike the parents, be a force in the physical sense of the term, then there is abundant provision in Nature for its indefinite replenishment. If, rather, it be a part or phase of that something which directs and determines the expenditure of force, then it is not subject to the laws of the latter, and there is no ground for inferring its exhaustibility. The limited vitality is an unproved and unprovable conjecture. The evolutive force, dying out in the using, is either the same conjecture repeated, or a misapplied analogy.

After all—apart from speculative analogies—the only evidences we possess which indicate a tendency in species to die out, are those to which Mr. Darwin has called attention. These are, first, the observed deterioration which results, at least in animals, from continued breeding in and in, which may possibly be resolvable into cumulative heritable disease; and, secondly, as already stated (p. 285), what may be termed the sedulous and elaborate pains everywhere taken in Nature to prevent close breeding—arrangements which are particularly prominent in plants, the greater number of which bear hermaphrodite blossoms. The importance of this may be inferred from the universality, variety, and practical perfection of the arrangements which secure the end; and the inference may fairly be drawn that this is the physiological import of sexes. It follows from this that there is a tendency, seemingly inherent, in species as in individuals, to die out; but that this tendency is counteracted or checked by sexual wider breeding, which is, on the whole, amply secured in Nature, and which in some way or other reenforces vitality to such an extent as to warrant Darwin's inference that "some unknown great good is derived from the union of individuals which have been kept distinct for many generations." Whether this reenforcement is a complete preventive of decrepitude in species, or only a palliative, is more than we can determine. If the latter, then existing species and their derivatives must perish in time, and the earth may be growing poorer in species, as M. Naudin supposes, through mere senility. If the former, then the earth, if not even growing richer, may be expected to hold its own, and extant species or their derivatives should last as long as the physical world lasts and affords favorable conditions. General analogies seem to favor the former view. Such facts as we possess, and the Darwinian hypothesis, favor the latter.

XIII

EVOLUTIONARY TELEOLOGY

When Cuvier spoke of the "combination of organs in such order that they may be in consistence with the part which the animal has to play in Nature," his opponent, Geoffroy St.-Hilaire, rejoined, "I know nothing of animals which have to play a part in Nature." The discussion was a notable one in its day. From that time to this, the reaction of morphology against "final causes" has not rarely gone to the extent of denying the need and the propriety of assuming ends in the study of animal and vegetable organizations. Especially in our day, when it became apparent that the actual use of an organ might not be the fundamental reason of its existence— that one and the same organ, morphologically considered, was modified in different cases to the most diverse uses, while intrinsically different organs subserved identical functions, and consequently that use was a fallacious and homology the surer guide to correct classification—it was not surprising that teleological ideas nearly disappeared from natural history. Probably it is still generally thought that the school of Cuvier and that of St.-Hilaire have neither common ground nor capability of reconcilement.

In a review of Darwin's volume on the "Fertilization of Orchids" * (too technical and too detailed for reproduction here), and later in a brief sketch of the character of his scientific work (art. IX, p. 234), we expressed our sense of the great gain to science from his having brought back teleology to natural history. In Darwinism, usefulness and purpose come to the front again as working principles of the first order; upon them, indeed, the whole system rests.

To most, this restoration of teleology has come from an unexpected quarter, and in an unwonted guise; so that the first look of it is by no means reassuring to the minds of those who cherish theistic views of Nature. Adaptations irresistibly suggesting purpose had their supreme application in natural theology. Being manifold, particular, and exquisite, and evidently inwrought into the whole system of the organic world, they were held to furnish irrefragable as well as independent proof of a personal designer, a divine originator of Nature. By a confusion of thought, now obvious, but at the time not unnatural, they were also regarded as proof of a direct execution of the contriver's purpose in the creation of each organ and organism, as it were, in the manner man contrives and puts together a machine—an idea which has been set up as the orthodox doctrine, but which to St. Augustine and other learned Christian fathers would have savored of heterodoxy.

In the doctrine of the origination of species through natural selection, these adaptations appear as the outcome rather than as the motive, as final results rather than final causes. Adaptation to use, although the very essence of Darwinism, is not a fixed and inflexible adaptation, realized once for all at the outset; it includes a long progression and succession of modifications, adjusting themselves to changing circumstances, under which they may be more and more diversified, specialized, and in a just sense perfected. Now, the question is, Does this involve the destruction or only the reconstruction of our consecrated ideas of teleology? Is it compatible with our seemingly inbore conception of Nature as an ordered system? Furthermore, and above all, can the Darwinian theory itself dispense with the idea of purpose, in the ordinary sense of the word, as tantamount to design?

From two opposing sides we hear the first two questions answered in the negative. And an affirmative response to the third is directly implied in the following citation:

"The word purpose has been used in a sense to which it is, perhaps, worth while to call attention. Adaptation of means to an end may be provided in two ways that we at present know of: by processes of natural selection, and by the agency of an intelligence in which an image or idea of the end preceded the use of the means. In both cases the existence of the adaptation is accounted for by the necessity or utility of the end. It seems to me convenient to use the word purpose as meaning generally the end to which certain means are adapted, both in these two cases and in any other that may hereafter become known, provided only that the adaptation is accounted for by the necessity or utility of the end. And there seems no objection to the use of the phrase 'final cause' in this wider sense, if it is to be kept at all. The word 'design' might then be kept for the special case of adaptation by an intelligence. And we may then say that, since the process of natural selection has been understood, purpose has ceased to suggest design to instructed people, except in cases where the agency of man is independently probable."—P.C.W., in the Contemporary Review for September, 1875, p. 657.

The distinction made by this anonymous writer is convenient and useful, and his statement clear. We propose to adopt this use of the terms purpose and design, and to examine the allegation. The latter comes to this: "Processes of natural selection" exclude "the agency of an intelligence in which the image or idea of the end precedes the use of the means;" and since the former have been understood "purpose has ceased to suggest design to instructed people, except in cases where the agency of man is independently probable." The maxim "L'homme propose, Dieu dispose," under this reading means that the former has the monopoly of design, while the latter accomplishes without designing. Man's works alone suggest design.

But it is clear to us that this monopoly is shared with certain beings of inferior grade. Granting that quite possibly the capture of flies for food by Dionaea and the sundews may be attributed to purpose apart from design (if it be practicable in the last resort to maintain this now convenient distinction), still their capture by a spider's-web, and by a swallow on the wing, can hardly "cease to suggest design to instructed people." And surely, in coming at his master's call, the dog fulfills his own design as well as that of his master; and so of other actions and constructions of brute animals.

Without doubt so acute a writer has a clear and sensible meaning; so we conclude that he regards brutes as automata, and was thinking of design as coextensive merely with general conceptions. Not concerning ourselves with the difficulty he may have in drawing a line between the simpler judgments and affections of man and those of the highest-endowed brutes, we subserve our immediate ends by remarking that the automatic theory would seem to be one which can least of all dispense with design, since, either in the literal or current sense of the word, undesigned automatism is, as near as may be, a contradiction in terms. As the automaton man constructs manifests the designs of its maker and mover, so the more efficient automata which man did not construct would not legitimately suggest less than human intelligence. And so all adaptations in the animal and vegetable world which irresistibly suggest purpose (in the sense now accepted) would also suggest design, and, under the law of parsimony, claim to be thus interpreted, unless some other hypothesis will better account for the facts. We will consider, presently, if any other does so.

We here claim only that some beings other than men design, and that the adaptations of means to ends in the structure of animals and plants, in so far as they carry the marks of purpose, carry also the implication of having been designed. Also, that the idea or hypothesis of a designing mind, as the author of Nature—however we came by it—having possession of the field, and being one which man, himself a designer, seemingly must needs form, cannot be rivaled except by some other equally adequate for explanation, or displaced except by showing the illegitimacy of the inference. As to the latter, is the common apprehension and sense of mankind in this regard well grounded? Can we rightly reason from our own intelligence and powers to a higher or a supreme intelligence ordering and shaping the system of Nature?

A very able and ingenious writer upon "The Evidences of Design in Nature," in the Westminster Review for July, 1875, maintains the negative. His article may be taken as the argument in support of the position assumed by "P.C.W.," in the Contemporary Review above cited. It opens with the admission that the orthodox view is the most simple and apparently convincing, has had for centuries the unhesitating assent of an immense majority of thinkers, and that the latest master-writer upon the subject disposed to reject it, namely, Mill, comes to the conclusion that, "in the present state of our knowledge, the adaptations in Nature afford a large balance of probability in favor of creation by intelligence." It proceeds to attack not so much the evidence in favor of design as the foundation upon which the whole doctrine rests, and closes with the prediction that sooner or later the superstructure must fall. And, truly, if his reasonings are legitimate, and his conclusions just, "Science has laid the axe to the tree."

"Given a set of marks which we look upon in human productions as unfailing indications of design," he asks, "is not the inference equally legitimate when we recognize these marks in Nature? To gaze on such a universe as this, to feel our hearts exult within us in the fullness of existence, and to offer in explanation of such beneficent provision no other word but Chance, seems as unthankful and iniquitous as it seems absurd. Chance produces nothing in the human sphere; nothing, at least, that can be relied upon for good. Design alone engenders harmony, consistency; and Chance not only never is the parent, but is constantly the enemy of these. How, then, can we suppose Chance to be the author of a system in which everything is as regular as clockwork? . . . The hypothesis of Chance is inadmissible."

There is, then, in Nature, an order; and, in "P.C.W.'s" sense of the word, a manifest purpose. Some sort of conception as to the cause of it is inevitable, that of design first and foremost. "Why"—the Westminster Reviewer repeats the question—"why, if the marks of utility and adaptation are conclusive in the works of man, should they not be considered equally conclusive in the works of Nature?" His answer appears to us more ingenious than sound. Because, referring to Paley's watch,—

"The watch-finder is not guided solely in his inference by marks of adaptation and utility; he would recognize design in half a watch, in a mere fragment of a watch, just as surely as in a whole time-keeper . . . Two cog-wheels, grasping each other, will be thought conclusive evidence of design, quite independently of any use attaching to them. And the inference, indeed, is perfectly correct; only it is an inference, not from a mark of design, properly so called, but from a mark of human workmanship . . . No more is needed for the watch-finder, since all the works of man are, at the same time, products of design; but a great deal more is requisite for us, who are called upon by Paley to recognize design in works in which this stamp, this label of human workmanship, is wanting. The mental operation required in the one case is radically different from that performed in the other; there is no parallel, and Paley's demonstration is totally irrelevant."[XIII-2] But, surely, all human doings are not "products of design;" many are contingent or accidental. And why not suppose that the finder of the watch, or of the watch-wheel, infers both design and human workmanship? The two are mutually exclusive only on the supposition that man alone is a designer, which is simply begging the question in discussion. If the watch-finder's attention had been arrested by a different object, such as a spider's web, he would have inferred both design and non-human workmanship. Of some objects he might be uncertain whether they were of human origin or not, with-out ever doubting they were designed, while of others this might remain doubtful. Nor is man's recognition of human workmanship, or of any other, dependent upon his comprehending how it was done, or what particular ends it subserves. Such considerations make it clear that "the label of human workmanship" is not the generic stamp from which man infers design. It seems equally clear that "the mental operation required in the one case" is not so radically or materially "different from that performed in the other" as this writer would have us suppose. The judgment respecting a spider's web, or a trap-door spider's dwelling, would be the very same in this regard if it preceded, as it occasionally might, all knowledge of whether the object met with were of human or animal origin. A dam across a stream, and the appearance of the stumps of trees which entered into its formation, would suggest design quite irrespective of and antecedent to the considerable knowledge or experience which would enable the beholder to decide whether this was the work of men or of beavers. Why, then, should the judgment that any particular structure is a designed work be thought illegitimate when attributed to a higher instead of a lower intelligence than that of man? It might, indeed, be so if the supposed observer had no conception of a power and intelligence superior to his own. But it would then be more than "irrelevant;" it would be impossible, except on the supposition that the phenomena would of themselves give rise to such an inference. That it is now possible to make the inference, and, indeed, hardly possible not to make it, is sufficient warrant of its relevancy.

It may, of course, be rejoined that, if this important factor is given, the inference yields no independent argument of a divine creator; and it may also be reasonably urged that the difference between things that are made under our observation and comprehension, and things that grow, but have originated beyond our comprehension, is too wide for a sure inference from the one to the other. But the present question involves neither of these. It is simply whether the argument for design from adaptations in Nature is relevant, not whether it is independent or sure. It is conceded that the argument is analogical, and the parallel incomplete. But the gist is in the points that are parallel or similar. Pulleys, valves, and suchlike elaborate mechanical adaptations, cannot differ greatly in meaning, wherever met with.

The opposing argument is repeated and passed in another form:

"The evidence of design afforded by the marks of adaptation in works of human competence is null and void in the case of creation itself . . . Nature is full of adaptations; but these are valueless to us as traces of design, unless we know something of the rival adaptations among which an intelligent being might have chosen. To assert that in Nature no such rival adaptations existed, and that in every case the useful function in question could be established by no other instrument but one, is simply to reason in a circle, since it is solely from what we find existing that our notions of possibility and impossibility are drawn. . . . We cannot imagine ourselves in the position of the Creator before his work began, nor examine the materials among which he had to choose, nor count the laws which limited his operations. Here all is dark, and the inference we draw from the seeming perfections of the existing instruments or means is a measure of nothing but our ignorance."

But the question is not about the perfection of these adaptations, or whether others might have been instituted in their place. It is simply whether observed adaptations of intricate sorts, admirably subserving uses, do or do not legitimately suggest to one designing mind that they are the product of some other. If so, no amount of ignorance, or even inconceivability, of the conditions and mode of production could affect the validity of the inference, nor could it be affected by any misunderstanding on our part as to what the particular use or function was; a statement which would have been deemed superfluous, except for the following:

"There is not an organ in our bodies but what has passed, and is still passing, through a series of different and often contradictory interpretations. Our lungs, for instance, were anciently conceived to be a kind of cooling apparatus, a refrigerator; at the close of the last century they were supposed to be a centre of combustion; and nowadays both these theories have been abandoned for a third . . . Have these changes modified in the slightest degree the supposed evidence of design?"

We have not the least idea why they should. So, also, of complicated processes, such as human digestion, being replaced by other and simpler ones in lower animals, or even in certain plants. If "we argue the necessity of every adaptation solely from the fact that it exists," and that "we cannot mutilate it grossly without injury to the function," we do not "announce triumphantly that digestion is impossible in any way but this," etc., but see equal wisdom and no impugnment of design in any number of simpler adaptations accomplishing equivalent purposes in lower animals.