Indian Pipe. (Monotropa uniflora). A saprophytic plant inhabiting rich woods in eastern North America. (Courtesy of Brooklyn Botanic Garden.)
The Partridge Berry (Mitchella repens), a trailing vine of northern forests. (Courtesy of Brooklyn Botanic Garden.)
Rafflesia. One of the Strangest Products of the Rain Forest. It consists only of a giant flower, the largest in the world, which is attached directly to the roots or stems of relatives of the grape, upon which it is parasitic. (After Kerner and Oliver. Courtesy of Brooklyn Botanic Garden.)
autophytes, literally solitary or self-providing plants, and this thrifty mode of life is called autophytic. But a few kinds of plants, actually many millions of individuals, have more devious ways of getting their food and provide strong contrast to their sturdier associates.
These baser modes of life appear to have been rather insidiously developed, as though there had been some hesitation at even the smallest departure from the normal. Of course we must not forget that plants, while living things, are never reasoning ones, and that good and evil and all other qualities that are ascribed to plants are perfectly foreign to them. Throughout this book, and in many others, the habits of plants are spoken of as base, for instance, or good. What is actually the fact is that nature works in truly marvelous ways, and to our reasoning faculties these adjustments seem clothed with attributes they do not really possess. But the description of them in the terms of our everyday speech, the translation of their behavior into the current conceptions of mankind, does so fix them in our minds that they cease to be “just plants,” and we no longer put their habits in the category of those interesting things that nearly everyone forgets.
One of the first signs of departure from the usual methods of getting food is the association of certain minute organisms at the roots upon which plants, otherwise autophytic, depend for aid in securing nourishment. This characteristic is fairly common, notably in all the plants of the pea family, such as peas, beans, locust trees, vetch, clover, and hundreds of others. If the roots of any of these be examined, it will be seen that attached to the smaller divisions of them are small tubercles from the size of a pinhead to a pea, depending on the kind. These tubercles or galls are caused by and infested with bacteria, the smallest of all plants. The bacteria have the extraordinary power of changing nitrogen into nitrates, which is the only form in which nitrogen can be absorbed by roots. Not only do they accomplish this, but excess nitrogen is stored in the roots by the same agency. It is this fact that has resulted in the planting of vetch and kindred plants for soil enrichment, as each year there is a residue of nitrogen left in their roots and by so much they add plant food to the soil. For hundreds of years farmers have done this, but only quite recently have we known why they did so. The occurrence of bacteria or microbes at the roots of plants is much more common than was formerly supposed to be the case, and many other plants than those of the pea family depend, at least in part, upon them in getting food from the soil. While not wholly autophytic, such plants do make some return for what they gain, as some of them at least pay dividends in extra nitrogen, and all of them provide opportunity for the bacteria to live. The latter play an important part in populating the soil, which is not the comparatively sterile thing it appears to be. Actually it is infested with organisms that play a mighty, if rather inconspicuous, part in the work of preparing the soil for plant growth. These organisms are so minute and the chemical nature of their work is so complicated that merely to mention their existence must suffice here. This close association of certain roots and bacteria, which, as we have seen, is of mutual advantage, is known as symbiosis. It is really only a kind of exchange, not unlike the storybook community that helped out by taking in each other’s washing. Unlike that community the association between the two works to the actual advantage of both, but the process is undeniably a step away from those wholly autophytic plants which live free and independent of such aid.
A much more gruesome habit of certain plants is their reliance for food only upon the dead. In the Indian pipe, some kinds of shinleaf, and in many other plants their roots and root hairs are changed or often nearly lacking, and we find them growing only on the dead bodies of other plants. One peculiarly repulsive characteristic of such plants is that they secrete at their roots a substance that hastens the decay of the dead, and, as if this were not rapid enough, there are associated with them certain kinds of minute fungus organisms that also speed up decomposition. Plants with this charming mode of life are known as saprophytes, literally sapros, rotten, and phytes, plants. “Rotten plants” they may be in their mode of life, but the pearly white stems and flowers of the Indian pipe have a certain ghostly charm, an almost statuesque beauty among the normal greenery of the gloomy dark woods in which they always grow. It is not without significance that Indian pipe bears no leaves, has none or almost none of the life-giving green coloring matter which we have seen to be the almost priceless possession of plants which lead a different, and perhaps a better life. The great bulk of saprophytes bear no leaves, and some only partially wedded to the habit appear to be midway between bearing normal green leaves and bearing none, or much reduced ones that are quite unlike the busy factories we know normal green leaves to be. Plants with this method of getting their food, must of course grow in places where dead and decaying vegetation is plentiful, and often as such soil is turned up there may be noted a peculiar dank odor, suggestive not only of its origin, but of the fact that these “rotten plants” make their home in it. Some of our most beautiful orchids grow in this fashion, but even there, in spite of flowers that for beauty of form are without rivals, the plants have no green coloring matter in their leaves, which are often reduced or even wanting altogether.
It might almost seem as if demoralization, so far as food habits are concerned, had reached its lowest point in these plants that literally rob the dead, but there are still lower depths to which certain plants have been reduced. This consists of robbing the living, and such plants are called parasites, a word perfectly familiar in other connections. Parasitic plants have no roots, but attach themselves to the roots of other plants, somewhat generously called hosts, from which they derive their food. The best known case is the common Christmas mistletoe, and the dodder (Figure 68), but there are hundreds of others. Nothing in all the realm of plant life so perfectly fits the action to the word as plants of this type, flourishing when the host flourishes, dying when it dies. Producing flowers and seeds, and often, by an irony of fate, perfectly green leaves, they are nevertheless the most debased of all plants in their mode of life.
These successive steps in the degradation of food habits, are not always the clean-cut things they might be inferred to be from the foregoing. There are many intermediate stages; it may even prove to be the case that some plants are wholly autophytic at certain stages of their life, and slip partially into more devious practices at other stages. The whole affair is not yet thoroughly understood and may well be the result of competition, as it is quite conceivable that if the getting of food in normal ways became difficult or impossible plants may have had to resort to other methods.
Some one has said that one day without water would make men liars, in two days they become thieves, and after the third or fourth day they would kill to get water. In the Army Records at Washington is a report of one of our expeditions, which in chasing Indians got lost in a desert, and in which the soldiers fought among themselves for even the most repulsive liquids. It hardly needs these gruesome examples, however, to confirm what everyone who has ever been mildly thirsty knows, that water is an essential for all animals, and that to be without it is to suffer torture. Air of the proper kind is just as important, and because its absence or impurity causes more sudden agony and a quicker death, the need of it is that much more acute. Plants rely even more upon these two essentials of life, and in getting them they behave in ways just as ruthless as do men who are suddenly deprived of either of them.
As we have already seen in “How Plants Get Their Food and Water from the Soil,” the water is the carrier of the food elements from the soil, but water as such does much more for the plant than act as a carrier. Osmotic pressure, a never-ending pump, keeps sending up a steady stream of water to the limits of its power. In everything except trees it seems fairly certain that this pressure is sufficient to drive water into the remotest leaves. It finally reaches these tiny rooms in the leaf about which we read in the account of Leaves as Factories. And just here a very curious thing happens. Each room is, as we have seen, a very busy place, crowded with all the necessary equipment to make sugar, and yet there is still room for water which is just as necessary as the other fittings; in fact so necessary is it that the whole interior of the room is bathed in water. This irrigation system works so well that the walls of the room literally bulge with the pressure of the water in them. If they did not—a condition known as turgor—the plant would at once wilt, and if no new supply came it would wither and die.
But water cannot stay in this condition of pressure and stagnation for even a brief period. That would be as if a leaf were like a toy balloon which, after inflation, had the entrance pinched and so remained inflated. And while we have all along spoken of factories for making sugar, and pressure pumps for forcing up food and water, it must never be forgotten that this marvelously adjusted mechanism is a living thing. Constantly growing, even producing their own means of falling in the autumn, leaves must be thought of as living machines, just as we are still more highly developed machines. In other words the accumulated water in the cells of the leaf must be removed, after it has served its use, and replaced by fresh supplies. The removal is carried on by its evaporation into the halls, or, in the more precise terms of our account of leaves as factories, into the intercellular spaces. It will be recalled that these are connected with the outside air through the pores or stoma. When the air outside is hot and dry it might easily suck out by evaporation all the water vapor in these intercellular spaces and wilting follow at once. This would actually happen if the guard cells, already mentioned, were not constantly on the job. They control the size of the opening just as certainly as a steam valve does, and maintain, with a few exceptions, just the proper amount of water loss not only to maintain turgor, but to see to it that transpiration, as this process is called, goes on rapidly enough to insure fresh supplies of water being sent to the leaf. The opening and closing of the stoma by the guard cells is a nicely balanced operation dependent upon root pressure, turgor, and atmospheric conditions. Guard cells have, because of this, been much studied in spite of the fact of their microscopic size. We now know that they allow greater openings during the night and reduce them during the day. When we reflect that the constant removal of water in the leaf, both as such, and as the only carrier of food supplies from the roots, depends in such large measure upon the functioning of these guard cells, then we come to some realization of their importance to the plant.
They do not always work unaided, for in many places the transpiration, even with their best efforts, would exceed the rise of water in the plant and death must follow if such a condition exists for long. This may be the case in certain bog plants, where, even with their roots in the water, they actually are in danger of drying out because the composition of bog water makes it partially unfit for most plants. And, again, in very open dry or windy places, such as deserts or the mountain tops above timber line, the actual supply of water may be insufficient. Many plants growing in such places have their leaves, particularly the under surfaces of them, clothed with various kinds of hairs. These may be quite velvety or cottony, but in any event, either by their texture or their color, they tend to reduce transpiration. An extreme case is a desert plant from Arizona where the whole leaf surface is covered with an ashy gray velvety coating, which, of course, absorbs less heat than a normal green leaf, and in addition there are much fewer pores through which transpiration could be carried on. In ever so many leaves nature has provided them with a thick coating of hairs in early spring, which they lose later in the summer. Shrubs and herbs, especially those that start earlier than the trees under which they grow, very often may be found with a dense woolly or silky covering in early spring. As the shade becomes denser and the need of the protection less, the wool or silk is shed, sometimes completely. Some of the most conspicuous cases of this are certain kinds of our common shadbush, which in April are covered with a beautiful grayish-white silky coat, but by August are practically the ordinary green color of other leaves. The hairy covering of leaves is well worth observation, as it may hide not a few facts about transpiration and, in some leaves, has had much to do with their preservation from grazing animals. Some, like the common mullein, are never touched, and may be found standing like sentinels in fields otherwise cropped short.
In many leaves there is conflict between those forces that result in the leaf getting the utmost possible exposure to light and those that prevent too rapid transpiration. On the one hand there is the absolute necessity for light, on the other the ever-present danger that the response of leaves to this necessity will result in a transpiration rate too rapid to be held in check by the guard cells. The compromise between these two forces, each pulling in opposite directions, gives to some leaves a series of movements that are among the most interesting things in nature. One of the most marked examples is the common wild lettuce, a weedy plant of our roadsides introduced from Europe. In bright sunlight the leaves are turned so that the edge of the blade faces upward, and the surface is thus protected from the direct rays of the sun, but during cloudy weather or in the shade the leaves turn into the ordinary position of most foliage leaves. It is difficult to avoid the inference that photosynthesis, which, as we have seen is an absolute necessity to the leaf, is in the wild lettuce retarded by transpiration, to avoid the too rapid rate of which the leaf is turned on edge. In this plant the leaf base, as though to be ready for whatever change transpiration or photosynthesis may demand, is so attached to the stem that such changes are made with the least possible delay or wrenching. In one of the many kinds of blue gum trees of Australia all the leaves turn one way in the light, and another in shade or on cloudy days. Ever so many plants have partial movements of their leaves, a good many of which are in response to these opposing demands, one pulling the leaf into the greatest possible light, the other holding it away from that condition. There are other movements of leaves, of parts of the flower, or even of the whole plant that are not so certainly the result of the conflict between light requirements and the necessity of conserving water supply. They will be considered presently.
While most plants are well provided with methods of losing water, so well provided in fact that in very hot or very long dry periods it is a common sight to see many plants literally panting for more water, there are some apparently more cautious individuals, who reverse this process. All throughout tropical America hundreds of relatives of the pineapple have their leaves so formed and arranged that they catch and hold considerable quantities of water. In one kind, called Hohenbergia, the long leaves are joined together toward their base into a water-tight funnel, which will hold a quart or two of water over a period of drought. In Africa the extraordinary traveler’s-tree, a giant herb growing twenty to thirty feet tall, has the overlapping leaf bases so arranged that they hold many gallons of water. And we have already seen how the giant cactus of our own Southwest will hold 125 gallons. The most remarkable case is the Ibervillea from the deserts of Arizona. In riding over this country one may find objects that look not unlike a burned pudding, about two feet in diameter and nearly as high. From the center comes a delicate stalk with the finest feathery foliage and tiny flowers. Of roots there appear to be almost none, and these curious objects, which are very hard and woody, might almost be taken for stones. But they are actually plants not distantly related to squash and pumpkin, and one of them collected years ago and brought into a museum behaved in quite the most thrifty fashion of any plant yet discovered. It was carefully cleaned and put in a museum case and locked up as a curiosity for the wondering public to gaze at. But suddenly, almost miraculously, it sent out its delicate growth which grew its appointed time and then withered. Imagine the astonishment of the curators of this museum to find it doing the same thing the next year, and the next. Finally after putting forth its shoot for five years it actually died and is now a peaceful museum specimen. No other such case of water storage is known, but thousands of plants have this remarkable ability to a less degree, all in response to conditions that would mean destruction to plants not so providently equipped.
This conservation of water on such a great scale offers striking contrast to the truly prodigal habits of certain plants that actually drip water, so charged are they with this precious liquid, and so little stress do their conditions of life put them under in this respect. Where water is plentiful and turgor maintained almost to the bursting point, evaporation in a moist or chilly atmosphere does not suck out water vapor fast enough. Sometimes, around the edges of the leaves of the common garden nasturtium, drops of water may be found, literally forced out as drops, rather than transpired as water vapor. This happens to a considerable number of plants, during the night when transpiration is laggard, and such drops are usually mistaken for dew. The latter is actually the condensation of moisture in the air upon the leaves of plants which cool down more rapidly than the air, and seldom due to the forcing out of drops of water from leaves, although in rare cases it may be. In tropical forests, where the humidity is very heavy and water supply from the roots copious, certain leaves leak water so fast and are so constructed that this excess is prevented from accumulating on the leaf. The pipal tree of India has long drip tips to its leaves that conduct the excess water from the blade to the end of the slender tip where it drips off. The advantage of these dripping points is obvious, for in regions so humid that water is forced out of the leaf, the coating of the leaf with this extra moisture would by that much retard transpiration. Dripping points, which in less exaggerated forms than in the pipal tree are common in many parts of the world, are thus of decided advantage.
Whether it be desirous to retain water or to lose it by gradual evaporation, or expel an excess of it, each species of plant has developed the apparatus to best preserve its individual life. While only the barest outline of these adjustments to the water requirements of plants has been given here, the details form an almost dramatic picture of struggle of the different kinds of plants for survival. The extremes are the desert plants on the one hand and those of the rain forests in the tropics on the other. The chapter on Plant Distribution will show how important these water requirements of plants have been in determining what grows on the earth to-day.
With carbon dioxide going in, oxygen, water vapor and, as we have seen, even liquid water coming out of the stoma of leaves, it might be surmised that these busy little pores and their guard cells had done work enough for the plant. And yet there is still one more act to play and the stoma have much to do with it. For this process of photosynthesis and the closely related one of supplying food and water to the leaf cannot go on without respiration, which is quite another thing. In plants respiration or breathing has no more to do with digestion than it does in man. Digestion in man is not unlike photosynthesis in plants, except that plants make food in the process while men destroy it. But plants must breathe just as we do, and, as we need oxygen to renew our vital processes, so do they. While respiration is a necessary part of plant activity it is not such an important part as photosynthesis, for which it is often mistaken. The thing to fix in our minds is that photosynthesis makes food, uses the sun’s energy and releases oxygen in the process, while respiration uses oxygen and might almost be likened to the oil of a machine—necessary but producing nothing.
In walking through the quiet cathedrallike stillness of a deep forest or over the fields and moors, perhaps our chief thought is how restful the scene is, and what a contrast the quiet, patient plants make to the darting insects or flitting birds that our walk disturbs. We found at the beginning of this book that ability to get about is one of the main differences between animals and plants. Like so many first thoughts, this is, however, only a half truth, for while most plants, seemingly by a kind of fatality, are anchored forever to the place of their birth, many of them do move certain parts of themselves and that quite regularly. While some of these movements have already been hinted at as a possible response to transpiration or too intense light, there are others where the advantage to the plant, if any, has yet to be demonstrated. These other movements, perhaps because their cause has never been discovered, seem the more mysterious as they certainly are more weird and interesting than almost any other of the curious things that plants do.
Perhaps the most difficult thing in the world is to keep an active growing child perfectly still for more than a few moments at a time. There seems to be some impelling force that makes young growing things in a constant state of restlessness, and it is perhaps not so extraordinary, after all, that practically all young plants are restless in the sense that they are never quite still. And, like many grown-up people who do not know what repose in their waking moments really means, there are a goodly number of plants that are restless until the day they die.
Charles Darwin, perhaps the greatest man that the last century produced, wrote a book in two volumes on these restless plants, and proved by a series of experiments illustrated by charts which the plants themselves drew for him, that there were perhaps no plants that do not move at least some part of themselves during the early stages of their career. While he never could explain the cause of these movements he left in that book an imperishable record of the amount and direction of these mysterious movements, which are almost to be likened to the growing pains of young children.
The tips or growing shoots of many plants will point in one direction in the early morning, a different way at noon and still a different one by nightfall. Hundreds of totally unrelated plants seem to have this habit of moving their tips through a definite cycle during each day and this restlessness does not appear to be of the slightest use to them. It cannot be response to the moving of the sun through the sky, for often the movement may be away from the direct sunshine, and sometimes the motion goes on in the dark, as experiments have proved.
It is hard to see the movement of the whole upper part of a plant, although it is well known that they do move in many cases. But in the tendrils the movement is often easy to observe and even to induce. Some of these slender aids to climbing plants, if they happen to be swinging freely in the air, do actually make slow circular movements, that even if they were designed for the purpose could not more perfectly accomplish their obvious intent, which is to catch the nearest favorable support. These circular movements are to the left in the hop, honeysuckle and many other plants, to the right in the climbing beans, morning-glory and some others. When the tendril reaches a support it almost immediately turns about it, in the same direction as its free movements through the air have been. It is thus this apparently aimless swinging of tendrils through space that determines whether the vine is going to twine to the right or left. The speed with which a tendril will take its first turns about a support is so comparatively rapid that, once the support is caught there is scarcely a chance of the vine being torn away by the wind or other agency as would surely happen if tendril movements were the leisurely things that some folks think they are. In the case of one Passion-flower vine, which are gorgeous climbers mostly from the tropics, the tendril made a complete turn in two minutes after it first touched a possible support. And there is a quite noticeable movement in thirty seconds if the tip of the tendril be ever so lightly touched. Teasing tendrils to see how much or how fast they will coil has resulted in some extraordinary cases of the “comeback” of some of them. Darwin irritated a tendril for a few moments and induced a partial coiling which straightened out when the object causing it was withdrawn. To see how long the plant would stand this sort of thing and still not be literally tired of coiling he succeeded in making the plant partially coil, and by withdrawing the incentive uncoil again, over twenty times in fifty-four hours. An impulse to coil of such persistence as this naturally results in vines forming the impenetrable thickets they do in many forests. It emphasizes how restless are the growing points of these climbers, and serves as a striking illustration of those gradual movements of many other plants that seem to have some relation to growth, but in a way not yet understood. For while it is an obvious advantage for the vine to swing its tendrils through the air this advantage has not yet been proved the cause of the swinging. In fact if all possible supports are removed the tendril will often coil anyway, a perfectly futile proceeding, that looks almost like disgust.
This general restlessness, which by the imaginative has been thought of as a mild protest by plants at their otherwise fixed position, is not so spectacular as that of certain other plants, notably the poplars. A flattened instead of a round leafstalk makes the leaves of these trees flutter in the lightest air and in a gale the tree is a mass of animated foliage. No use has ever been found for this curious habit and it is not certain that it is of the least advantage to the tree. If anything, the constant movement may have the decided disadvantage of increasing transpiration.
In our common wood sorrel the leaflets on cloudy days or during the night regularly “go to sleep.” That is, they are folded at such times, rather than spread out in the ordinary way. These sleep movements may have something to do with transpiration, but whether or not this is true they are very regular and in certain plants the habit is remarkably and rather mysteriously uniform. Why, for instance, do the leaflets of these wood sorrels, the beans, lupine, locust tree and licorice plant, always fold downward while the clovers, vetch, peas, and bird’s-foot trefoil are always folded upward? Such movements and their direction are among the unsolved problems of botany, and merely to know of them or observe them leads us nowhere as to their true inwardness.
But quite apart from these merely restless plants, and there are thousands of different kinds which are known to move slightly, at least during their young stages, are a few more decidedly active ones that are seemingly irritable. At least they show peculiar movements if touched, and at night. One of the best known is the sensitive plant from tropical America. Its twice compound leaf is composed of many tiny leaflets which upon the slightest touch close up and apparently wither on their stalk at once. In five seconds after the leaf is touched it will appear like a wilted wreck. If the jar is sharp enough the whole plant will droop, and the response to a sudden jar is almost electrically quick in its action. And yet all this sudden wilting, actually caused by a quick loss of turgor, is slowly repaired and the plant carries on quite normally again until another shock renews its irritable response. This plant does the same thing gradually during the night, except that the leaflets recover their normal position only with the rise of the sun.
From India comes the most remarkable of all plants so far as movements are concerned. For in the telegraph plant the movements are so regular and long continued that irritability might almost be said to be continuous. The plant is a low shrub or herb with compound leaves, and the terminal leaflet, which is much larger than its neighbors on either side of their common stalk, performs a motion that describes with its tip an irregular oval or ellipse. But the movement is not steady; it goes by a series of slight but perfectly distinct jerks. It takes about two minutes for the leaf to complete its cycle, and it is this jerky movement that has given the plant its name. During the night its leaflets stop this apparently quite useless performance, the cause of which is quite unknown. It is often grown in greenhouse collections where its strange movements may be seen on any sunny day.
Many other cases of the restlessness or irritability of plants could be given, and nothing has been said here of the curious movements of some insectivorous plants as they have already been mentioned. The very considerable movements of certain flower and fruit organs will also be considered elsewhere.
It cannot have escaped the thoughtful reader that all of this chapter on plant behavior has dealt with those functions of plants in which roots, stems, or leaves play the chief part. These purely vegetative actions of plants, what might almost be called their bread and butter activities, would never lead to perpetuating their kind. For while all of these functions are necessary, except certain apparently wayward movements which still remain unexplained, they are in a sense only the preparation for an infinitely more important act, the reproduction of their kind. What the poetic have called the love of the flowers, or in more prosaic but perhaps more truthful words the fertilization, pregnancy, and birth of the new race, will be considered in a separate chapter. No other act of the plant world is so interesting as the mechanism of reproduction, the almost endless devices for securing it, and the ingenuity of nature in seeing to it that there are no flukes.
THERE is perhaps no device of nature that more perfectly accomplishes its purpose than the one with which all living things are endowed—the instinct for the renewal of life. In man the dawn of the mating instinct has ever been the theme of poets, and some of its manifestations are the despair of ascetics. Through it some of the noblest of man’s emotions have arisen, and because of its perversion our daily newspapers chronicle the basest and most sordid tragedies.
But whether noble or ignoble, this instinct for mating is, in its simplest terms, only a provision of nature that all life contains within itself the means of renewing life. Without this, life, so far as we know it, would end with the present generation. Perhaps our understanding of this decree of an all-wise nature to increase and multiply will be heightened by looking at it not only from its familiar manifestations in man, but more broadly. Seen from this broader viewpoint, it is the inherent legacy of all living things from the dawn of life on the earth down to the present. Even the simplest one-celled organisms have the faculty of increasing. In all plants, both the flowerless ones and those producing flowers, the process is carried to a perfection almost unbelievable in its intricacy and in provisions against its failure. From the matings of flowers much may be gleaned; even man himself can learn from them the capacity for sacrifice, the sinking of individual aims and pleasures in the greater scheme of conforming to that necessity for renewal of the race upon which all progress must be based.
The equipment which different flowers have developed for this purpose, their almost uncanny devices to make certain that only the distant and foreign male can ever impregnate the female, the enormous wastage of both unfertilized females and males that will never become fathers, and the overwhelming effectiveness of it all, in spite of this prodigality—these manifestations of the production of young in the plant world will take up the rest of this chapter. All the first part will tell of this process in flowering plants, while the second shows how flowerless plants accomplish the same end in more secret ways. Finally, in a brief third part, we shall see how, without mating of the sexes, nature has still one other way to see to it that there is a constant supply of young.
We have already made clear that all plants are divided upon the basis of whether they bear flowers and their mating goes on before the world, or whether they bear none and the process is accomplished in more secret ways. Because flowers are so much better known, and it is simpler to see how the act is consummated in them than in the cryptogamous plants, we shall first consider the phanerogams or flowering plants, and in the second section of this chapter the cryptogams or flowerless plants.
In the first chapter, under the section devoted to flowers, we found that the stamens are the male and the pistils the female organs of reproduction. As the period for mating draws near there is developed in the anther, which is the enlarged tip of the stamen, a fine, usually yellow, powder known as pollen. This matures in the anther, and when ripe is discharged from tiny pores.
Pollen is made up of individual pollen grains, which are very often stuck together so that we see only the mass, not the individual pollen grain. Sometimes the pollen is not sticky, as in the case of pine trees or in the ragweed—a fertile cause of hay fever. In these, and hundreds of other plants, the wind will blow great clouds of pollen through the air. When we stop to consider that a single, or at most a very few pollen grains are all that are necessary—in fact, are all that can be of real service—the enormous wastage of the male fertilizing substance, in order that mating be secured, gives us some idea of how prodigal is nature in this supreme function.
The pistil, or female organ of reproduction, is more cautious in the expenditure of its resources. As we have seen, it is composed of a swollen base, the ovary, a slender shank, the style, and a swollen or branched tip, the stigma. In some plants the ovary is divided into several compartments or cells, each with one or more ovules, which are only immature or unfertilized seeds, often very tiny, but usually quite easily seen if the ovary is cut open. It is the entrance of the pollen grain into this ovule that consummates the act of fertilization. As the ovule is carefully secreted within the ovary of the flower, and as the male fertilizing stuff or pollen is found only on the anther, it is obvious that some method of bringing the two together must be provided for.
In some plants this is accomplished by the anthers being just above the stigma, and when the pollen is ripe and the ovule ready, the stigma is found to be covered with a sticky substance. As the falling pollen grains touch the stigma, they are caught in this sticky substance just as surely as flies are caught once they touch a fly paper. But just here one of the most wonderful processes of nature begins. The pollen grain begins, slowly at first, to grow, and in the act it penetrates the outer coat of the stigma with a minute pollen tube. This slender threadlike tube, carrying with it the male germ, grows straight down through the stigma, into the narrowed style, and through this to the ovule. Once the pollen is caught on the stigma, nothing is so sure of fulfillment as that this male fertilizing stuff will ultimately reach the ovule. For the hitherto virgin ovule this impregnation starts a new phase in life. It means the beginning of the end, but in the process fruit and seed will be developed, and the young bride, already a mother, has triumphantly accomplished that for which she exists.
If fertilization of all flowers were as simple as this, there would be no need of what follows, but actually in surprisingly few plants are the stamens and pistils so arranged, the ripening of the pollen and readiness of the ovule for impregnation so timed that the act can be accomplished in such direct fashion. For it is quite obvious that in flowers in which the whole drama of mating goes on within the petals, without the interference or help of any outside agency, the result will be a crop of young who know no other characters than those of the parents, and have nothing to look forward to but a closely inbreeding progeny, very little, if at all different from themselves. In other words, such plants are pure bred, they lack the usually obvious virility that comes from crossing the male of one plant with the female of another. There are so many devices to prevent self-fertilization in flowers, so marvelous are the contrivances to see to it that only cross-fertilization can be effective, and, finally, the experience of breeders that strength and virility often or usually result from impregnating the ovules of one plant with the pollen of another, that we are forced to the conclusion that absolute purity in the sexual relations of flowers is rare indeed. It occurs, without peradventure of a doubt, only in those flowers whose petals never open and where fertilization is consummated, if not in private, at any rate without external help. In many violets the showy violet blossoms are often nearly infertile, while down near the ground are inconspicuous flowers which never open, but within which fertilization is so successful that the crop of seeds is far more plentiful than in the more showy ones that most people think are the only flowers ever borne by violets. These flowers that never open, or at any rate open so slightly that their sexual processes are modestly completed without intrusion, are known as cleistogamous flowers (Figure 69). They have been found in a few plants, but overwhelmingly the greater number of flowers not only do, but must, rely on some outside agency to insure fertilization.
Certain structural features of flowers have been so developed that fertilization of the ovary by the pollen of the same flower is impossible. The commonest case is in those flowers where the stamens are shorter than the pistils, as they always are in the common snowdrop, hyacinth, the sassafras tree, and in hundreds of others. There can be no consummation of the reproductive process in such flowers without some outside aid. More futile still without this aid are those flowers where the stamens are well above the pistils, but the time of maturing in both differs by a few days or even hours. Nothing could be more helpless than the pistil under these circumstances, for if its instinct for maternity were ever so strong, it would be doomed to barren sterility by the premature development of the males. Sometimes, too, the female is prematurely ripe for impregnation, and the stamens lag behind a day or two. Her time passes and with it her only chance of fertilization—by her own haste she has rendered impotent the now useless pollen which appears doomed to fall aimlessly upon the unreceptive stigma.
But, perhaps, the most hopeless of all is the well-known partridge berry, whose red berries are common in the woods during August and September. This seems as though it fought off any chance of securing a mate by a flower structure and behavior that would certainly so result if some way out of the difficulty were not at hand. The partridge berry bears two kinds of flowers that outwardly look much alike, but whose sexual organs differ in this way: in some flowers the stamens are all shorter than the pistil, and in others the pistil is much outtopped by the stamens. The extraordinary feature of it is not so much this structural difference, however, but the fact that pollen from the short-stamened flower is useful only to its neighboring short-styled relative, while the pollen from this long-stamened but short-styled neighbor is nearly useless where it is found and really useful only on the long-styled plant. By this device, but again only with outside aid, this plant does not prevent maternity, but increases its chances of being fruitful, for, as we have already seen, cross-fertilization appears to be the rule rather than the exception, and the partridge berry not only needs it, but can exist only when its offspring are the result of such crosses.
In all those plants that bear the different sexes in different flowers on the same plant, as in the hickory, or even on different plants, as in the willow, there must, of course, be some method arranged for cross-fertilization or they would promptly die out. So general is this cross-fertilization, so much a part of the economy of nature does it appear to be, that we can only think that there must be in the production of this vast horde of the cross-fertilized some advantage. Besides securing greater virility, which almost certainly results from this promiscuity, greater variability is promoted. If virility is the result, the price paid for it is tremendous, for the hindrances to self-fertilization are so many and so effective that most flowers would inevitably die as perfectly pure but ineffectual virgins if that fatality were not prevented. How they are saved from such a sterile fate, how they finally secure a mate by devices that outshine the most bewitching tricks of the daughters of Eve, is one of the most fascinating stories in all the history of the plant world.
For, of course, flowers do secure a mate, and they are aided in this enterprise by the most formidable array of helpers, one might almost call them conspirators. The chief of these are insects, thousands of different kinds of which are constant flower visitors. Some of the smaller birds, and even snails, also help flowers to meet their mates. The wind, too, bears pollen through the air to some expectant bride-to-be. And, finally, in the water, by a series of acts the like of which no one could improve for cunning, the cross-fertilization of certain aquatic plants is consummated. It would take a book larger than the present one to give even the briefest account of how these different aids to maternity do their work and how the flower responds to this help. As that is quite out of the question, only some of the best-known examples of cross-fertilization will be given, and these will be grouped according to what agency the flower is indebted for its chance of doing that for which it is created.
On any summer day, especially when the sun is shining brightly, we may see bees and butterflies flitting from flower to flower, busy as the proverbial bee. We already know enough about nature’s ways of doing things to be certain that these, and hundreds of other kinds of insects, do not come for nothing, and that the flower must have something to offer. Bees, especially, are thrifty creatures whose business demands exacting and prolonged toil. They would not waste five seconds upon idle flower calling if the blossoms did not yield a rich store. And thousands of flowers do yield the sweetest and richest kinds of stores of nectar or honey, which is, in fact, by the help of insects who alone can extract it, our sole source of honey. Many flowers which produce no nectar do have such plentiful stocks of pollen that the bees come for that alone. In the peony, for instance, over three million pollen grains are produced in each flower, only a minute fraction of which can ever fertilize an ovule. All the rest would be wasted were not pollen in itself a particularly nutritious food for young bees, and consequently much sought after by the careful bee mothers. They are the only insects that feed their young on pollen, or beebread, as it is called by the beekeepers, so that the enormous overproduction of this male fertilizing agent, from the point of view of the flower, is a decided attraction, one might almost call it a trap, to insure constant visitations from bees. For they are perhaps the most useful of all insects in the great game of securing cross-fertilization, as we shall presently see. Many other kinds of adult insects eat pollen directly and so add to the number of insect visitors.
No one has ever been able to explain the beautiful coloring of flowers, except that it serves as an attraction for insects and small birds. Like the honey or nectar, it seems to play no real part in the home economy of the flower, to be of not the least use otherwise. While honey and the gorgeous colors of flowers are a delight to man, that would be no sufficient reason for the ability to produce them. Both of these attributes of flowers, as attractions for insect visitors, are, however, so absolutely essential to cross-fertilization that we must think of them as having grown up out of that demand. As we shall see a little later, even the structure of some insects has been modified so that they can reach the nectar or pollen only by automatically doing for the flower what it cannot do itself.
While color of flowers seems as though it were attraction enough, it is very likely that their fragrance or perfume is still more seductive in its power of luring insect visitors and repelling useless ones. Poets have called this perfume the soul of the flower, and in its almost intangible beauty it might well be so called were it not for the fact that it appears to be of not the slightest use, except as a lure. In all the equipment of seduction there is none like this fragrance of flowers for attracting insects.
Flowers, then, have things to offer to insects which the latter need. Nectar and pollen are the chief, and where these merely bread and butter objects are not enough, or sometimes in addition to them, the flower is dressed out in gorgeous colors or perfumed with a fragrance beyond the dreams of the fairest bride. What insects do to complete the fertilization of such a legion of beauties makes up the romance of the flowers. Perhaps not even in man himself is this creation of new life so surrounded with beautiful ideas. Also, as in man, it sometimes is bound up with an almost fiendish cruelty and cunning. Some of these visitors and what they do, but unfortunately only a very few, can be mentioned here. They must serve as types or examples of what is going on all about us on any summer day.
The common blue columbine, much grown in gardens for its beautiful blossoms, always has the flowers hanging upside down, a habit that admirably serves to keep its pollen from rain. The opening and closing of many flowers in cloudy weather, or at night, may be for the same reason. Everyone knows the five blue spurs into which the petals of columbine are produced. At the very end of each spur, which is always curved, the flower secretes a considerable quantity of honey. This, one of the greatest attractions to bees, leads inevitably to a visit from one. The bee, in order to reach the honey, hangs on to the inverted flowers, clutching the base of the spur with its foreleg, and further securing itself by the mid or hind legs, which grasp the slender column into which, in the columbine, the stamens and pistils are crowded. In its anxiety to reach the honey the bee pokes its head as far into the spur as possible, but it gets in only a fraction of the full length of the tube. To reach the honey it extends its sucking apparatus, which is a complicated mechanism for this purpose on the head of nearly all insects, and which will hereafter be called by its true name of proboscis. It happens that bees can easily bend the proboscis downward or toward their own body, but only with considerable difficulty can they bend it in the opposite way. And yet the honey in the curved tip of the columbine can only be reached by curving the proboscis to fit the tube, and in this process the bee’s body for nearly half its length is forced to touch the anthers. While these are close to the stigma, they produce pollen only on their outside surface, where it is, of course, scarcely likely to reach the stigma, but must be brushed off by the contortions of the bee’s body in reaching the honey. The hairy body of the bee, coated with pollen, goes next to perhaps an older flower of the columbine. Heedless of any change in the flower the bee goes straight for the honey in one of the spurs, again catches hold of the only available support in the center of the flower. But this time, instead of brushing pollen off the exposed anthers, it brushes it off its own body to the stigmas, which, at a slightly later stage than in the one just described, are branched and perfectly adapted for collecting the pollen with which the bepowdered bee can hardly avoid dusting them. Cross-fertilization is of course assured, but it seems a precarious business at best, as the number of bees with a proboscis long enough to do the work is limited. The columbine, by a kind of uncanny foresight, is so constructed that bees or other insects that try to reach the honey and are not provided with a sufficiently long proboscis, nevertheless in further attempts upon other flowers, inevitably cross-fertilize them without reaping their reward. One or two kinds of bees, as though in retaliation for this subterfuge of the columbine, make short work of the honey by biting a hole in the spur and forthwith sucking out the honey without so much as touching anther, pollen, or stigma. The reply of the columbine to this ravaging of its chief attraction is that finally, as a last resort, and by a new movement of its reproductive organs, it is self-fertilized. Here the shape of the flower, the original position of the pistil and anthers, the exposure of pollen only in such a direction that, while a chance of cross-fertilization still exists, it can hardly ever fertilize its own stigmas, all point to cross-fertilization as the plant’s greatest requirement. And yet failing this, it falls back on self-fertilization rather than endure barren sterility.