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 a.s.sociated 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 s.p.a.ce 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 acc.u.mulate 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 princ.i.p.al 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 a.s.sured 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 quant.i.ty 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, alb.u.men, 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 pa.s.ses 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 circ.u.mstances, 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 r.e.t.a.r.ds 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 compa.s.s, 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 Pa.s.siflora 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 pa.s.ses 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 s.e.xUAL 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 a.s.signed 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, s.e.xually 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 const.i.tuted 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 circ.u.mstances. 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 a.n.a.logy 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 ma.s.s 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-s.e.xually-propagated varieties are endowed with the same power of unlimited duration that is possessed by varieties and species propagated s.e.xually--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 succ.u.mbing, nor to prove that a parasitic fungus (Peronospora infestans) is what is the matter with potatoes. For how else would const.i.tutional 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-s.e.xually 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 G.o.ds 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 a.s.serted 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 s.e.xual 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 s.e.xual 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 s.e.xual reproduction.

The conclusion of the matter, from the scientific point of view, is, that s.e.xually-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-s.e.xually 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