leaves, and persist for a couple of years, when they pass away, leaving slender, round, brown stems. In our native Broom (Sarothamnus scoparius) a similar modification may be observed, though of less degree. Sometimes stem-structures assume a very leaf-like form, as in the Butcher’s Broom (Ruscus aculeatus), where the ultimate branches are ovate and quite flat, and might be taken for true leaves but for the fact that they bear on their surface flowers, and subsequently berries. The leaves themselves are in this plant reduced to minute scales, and from their axils these flattened branches spring. In fact, where leaf reduction takes place, the process of assimilation is often shared in varying degree by the leaves, the stipules, and the stems. Among our native plants, as, for instance, in the Leguminosæ and Rosaceæ, the reader may find for himself many interesting examples for examination.
But the large majority of the Seed Plants bear well-developed leaves, to which the process of assimilation is practically confined.
Leaves vary surprisingly in size, shape, and arrangement, features which are closely related to the characters of the stems which bear them, the object being the most advantageous display of the chlorophyll in relation to the light-supply. In general they naturally take the form of a broad thin blade, protected as may be necessary against extremes of weather, and guarded against the obvious danger of being dried up by a thin waterproof covering or cuticle outside the epidermal layer of cells. In leaves we find the same beauty of mechanical construction as is seen in stems. The problem is again that of securing maximum efficiency with minimum expenditure of material. To give as great a surface as possible, the leaves are as broad and thin as is consistent with safety, the question of damage by wind being an important controlling factor. The veins, or vascular bundles, act efficiently as strengtheners of the thin surface; to prevent tearing at the leaf-edges the veins are often looped along the margin; while in indented leaves the extremities of the indentations are strengthened with special tissue. When one surface of the leaf faces the sky, as in most cases it does, this surface is strengthened against the weather, and the stomata are arranged mostly on the lower surface. Where occasionally the leaves hang normally in a vertical position, as do the mature leaves of the Gum Trees (Eucalyptus), both sides are protected, and the stomata are borne on the two faces equally. In the Water Lily, again, whose leaves float, the upper face, which alone is exposed to the air, bears the stomata, which are present in unusual numbers—nearly 300,000 to the square inch; the leaf surface is toughened to resist rain and wind, and waxy to prevent water from lying on it and so interfering with transpiration. The presence or absence of a leaf-stalk, again, is often clearly related to the light question. In the Water Lilies the continued lengthening of the elongated petiole causes the older leaves to float clear outside of the younger ones. In many biennial herbs, where food is stored up during the first season in preparation for the flowering effort in the second, a similar arrangement prevails—note the leaf-rosettes displayed by Spear Thistle (Carduus lanceolatus) and Herb Robert (Geranium Robertianum), as also especially in winter by perennials like the Dandelion (Taraxacum officinale) and Ribwort (Plantago lanceolata). Where stems spread horizontally, as the lower branches of trees, the leaves are arranged more or less in one
plane, in such a manner that overlapping is reduced to a minimum (Fig. 21). This is well seen in horizontal branches of the Elm and other familiar trees. In the plant chosen for illustration (Azara microphylla, a Chilian shrub), an interesting arrangement obtains. One of the pair of stipules which subtends each leaf is itself leaf-like, and stands at an angle, so that a mosaic is formed of true leaves (the larger ones) and stipules (the smaller alternating ones). On all stems the leaves are arranged not at haphazard, but according to definite rules. Sometimes they
are grouped in circles (whorls) at certain points of the stem, as in the Bedstraws; often in opposite pairs, arranged criss-cross, as in the Sycamore; most frequently in a series of spirals. The result in all cases is the same—it allows of as great an interval as possible between any leaf and the one immediately below or above it, and gives to all an equal share of light. The indenting of leaves, as in the Sycamore, or their division into separate segments, as in the Ash and Horse Chestnut, is of undoubted advantage as allowing light to pass through to lower layers of leaves; it also materially diminishes the danger arising from excessive wind-pressure. In the former case there is often a wide space between the divisions of the leaf; but where this is not required, the parts of the leaf fit closely together, to secure a maximum of surface. A particularly pretty example is seen in the Chilian shrub Weinmannia trichosperma (Fig. 22). Here, to avoid the loss of the area between the leaflets, the mid rib steps in, developing triangular wings which fill the spaces. It might be objected that the plant might have saved itself much trouble by producing, while it was about it, a simple undivided leaf covering the whole area. It is difficult to answer such suggestions. Probably the present form of the leaf best meets the conditions of wind, rain, and light under which it lives. Possibly its present form is bound up with its ancestral history. “It must be acknowledged,” says D. H. Scott, “that nothing is more difficult than to find out why one plant equips itself for the struggle with one device and another attains the same end in quite a different way.”
During cold and tempestuous weather the presence of leaves may be a danger to the plant rather than a help; and where seasonal variations are such that strongly contrasted periods of favourable and unfavourable weather occur, such as the summer and winter of our own climate, many plants have adopted the device of shedding all their leaves: this is especially characteristic of the largest plants (the trees), which would naturally suffer most from unfavourable weather. The fall of the leaf is accomplished by means of the formation of a transverse layer of corky tissue across the base of the leaf-stalk, combined with a weakening of the layer of cells immediately above. Prior to the perfecting of these arrangements for dropping the leaf, all the useful materials in it are withdrawn down the stem, so that only an empty skeleton is shed; the scar that remains is not an open wound, but is well protected by the corky layer before mentioned.
Stipules and bracts need not delay us in this sketchy survey of plant organs. They are leaves, generally of rather small size, placed, the former one on either side of the point where a leaf-stalk emerges from the stem, the latter singly below a flower; they are present in some plants, absent from others. They function in the same way as ordinary leaves, and in the earlier stages of growth are of use protectively. Occasionally the stipules exceed or even replace the leaves, as in the native Lathyrus Aphaca, where the leaf is reduced to a tendril, and the pairs of broad “leaves” are really the stipules. The bracts, in their turn, sometimes take on the “advertisement” function of the petals, as we have already seen (p. 87) in the case of certain Euphorbias.
The leaves of water plants offer several points of interest. Where they are entirely submerged, and, protected against the drying influence of wind and sun, they are of filmy texture. Broad blades are seldom met with, the leaves being usually either finely dissected or strap-shaped. The floating leaf, on the contrary, as already described in the Water Lily, is strongly built up, to withstand wave action and rain; it is usually broad and entire, which simplifies the
problem of avoiding submergence; and the stomata are confined to the upper side, which alone is in contact with the atmosphere. Those water plants which raise their leaves into the air, on the other hand, have leaves of a variety of shapes, which in most respects approach those of land plants. An interesting progression of leaves illustrating all three stages may be watched in spring in the Arrow-head (Sagittaria sagittifolia). The first leaves produced are entirely submerged, and conform to the usual ribbon shape and delicate texture. Those which follow float on the surface. In them the lower part is contracted into a flaccid winged petiole, the upper part being expanded into an oblong floating blade with a waxy surface to keep the leaf dry on the upper side. These in turn give way to the characteristic aerial arrow-shaped leaves of summer, which approach in character the leaves of land plants, and are borne on stout, stiff petioles capable of resisting wind and wave.
Coming now to FLOWERS, it is possible here to refer only to a few macroscopic or “naked-eye” characters and modifications; the full study of the flower and its essential functions being a matter for the laboratory and the high-power microscope, as very minute structures are involved. As briefly described in Chapter IV., flowers are groups of modified leaves arranged mostly very close together at the ends of branches, the tip of the shoot being often expanded into a receptacle (very well seen in the Compositæ—e.g., Dandelion) for the accommodation of the crowded floral leaves. Just as the foliage leaves have become modified to carry on to the best advantage the process of assimilation, so the different series of floral leaves are specially adapted to their several functions. The sepals, which compose the calyx, having usually a protective rôle, in most cases enclose the young flower with a tough envelope; they usually retain their primitive green colour, and take part in the process of assimilation. They may drop off as the flower opens (e.g., Poppy), or wither as the petals wither, or remain fresh until the fruit is ripe. Sometimes, as in many Ranunculaceæ (compare Anemone, Caltha, Helleborus), they take on the advertising rôle usually assigned to the petals, being large and coloured, while the petals themselves are minute. In the Monocotyledons they usually join with the petals in adorning the flower. The next whorl, lying inside (that is, above) the sepals, is formed of petals, constituting the corolla. The connection of colour and form of petals with the visits of insects, and their relative insignificance in wind-pollinated flowers, has already been referred to (p. 81). The marvellous variety of colour and form observable in the corolla has for its main object the attracting of insects to the flower. The petals have departed much farther from the ordinary leaf-form than the sepals. They assume brilliant hues of every tint, the pigment being due either to colouring matter dissolved in the cell-sap (pinks and blues) or to small coloured solid bodies (chromoplasts) contained in the cells (reds and yellows). Chlorophyll being absent, the coloured petals do not assist assimilation: they are purely advertisements, though incidentally they often fulfil a useful protective rôle for the important organs which they surround. In this latter connection their sensitiveness to changes of light and temperature, which causes them to close in dark or cold weather, is a very familiar phenomenon; as is also the excellent protection which they provide in flowers such as those of the Labiatæ, where, fused together into a tube, they form a kind of cave in which the stamens and pistil nestle securely.
An exceptional use of petals, where indeed they are used for the purposes of advertisement, but to secure the dispersal not of the pollen, but of the seeds, is illustrated in Fig. 24. In the genus Coriaria the staminate and pistillate organs are borne on separate flowers. The flowers of both kinds are small and inconspicuous. But in the “female” flowers the petals persist after flowering, and, becoming fleshy and comparatively large, enclose the seed in a pulpy berry-like envelope, which no doubt serves the same purpose as a true berry in securing seed-dispersal by being devoured by birds. In C. terminalis, which comes from the Himalayas, the “ripe” corolla is bright orange; in C. japonica, from Japan, it is at first coral-red, and when mature velvet-black.
The stamens, which form the next ring (sometimes a double ring or a close spiral), are much less leaf-like than the sepals or petals, yet there can be no doubt that they are descended from leaf-shaped organs; this is especially clear from the study of certain primitive fossil types, in which the corresponding organs which bear the pollen are actually leaf-like. In most of the present-day Seed Plants the stamens conform to a uniform type—a slender stalk (filament) bearing a head (anther) containing four chambers, in which are produced pollen grains, which escape when the flower is mature by the splitting of the enclosing walls. The ways in which the pollen is then conveyed to the pistil of other flowers have been referred to briefly on a previous page (p. 82). The stamens in many flowers are few, and their number usually bears a relation to the number of the other floral parts; in other flowers, for instance Rose and St. John’s wort (Hypericum), they are of large and indefinite number. The peculiar arrangement of the pollen in Orchids has been already noted (p. 94).
The final ring of modified leaves in our typical flower constitutes the pistil, formed of one or many carpels, the essential structure of which has been touched on already (p. 82). In the present place it is desired only to point out some of the leading modifications which the pistil undergoes, so that its structure as seen by the naked eye may be understood. In the simpler forms of carpel, the affinity to leaves is still evident, though in forms of pistil made up of a number of carpels this may be very difficult to trace. With the Pea, for instance, we may begin, as presenting a very simple example. Take an oblong leaf like that of a Laurel, and fold it down the mid rib till the two edges are in contact. There is our pea-pod complete. The young seeds, or ovules, are borne in a row along the mid rib, a very usual arrangement. Examine next the young fruit of a Columbine (Aquilegia). Here there is a group of five separate erect carpels, but each is essentially like a pea-pod in structure. Compare the fruit of a Saxifrage. This clearly consists of two carpels which are grown together save at the tips, where the two styles stand out like little horns. From this we may go on to other pistils in which several carpels are completely fused together. Next, the compact body thus formed may be sunk down in the expanded top of the stem (the receptacle). Or the other parts of the flower—sepals, petals, stamens—may in their lower part be fused with the walls of the pistil, and may thus appear to spring from the top of it. In such cases the structure of the flower may easily be wrongly interpreted, and reference to a work on systematic botany is necessary if pitfalls are to be avoided. It is indeed to be noted that in flowers, as in other parts of plants, complicated structure or multiplication of parts is not necessarily an indication of advanced evolution; on the contrary, it is often indicative of a primitive condition. Just as in machinery or in organized human effort simplification often accompanies improvement, so it is with plant structures. Many of the more primitive types of flowers, such as Buttercups or Water Lilies, have a multitude of petals or stamens or carpels, while in many of the most specialized, such as Composites or Campanulas, the number of parts is much reduced. The primitive wind-pollinated flowers produce large quantities of pollen; in those which have adopted the improved method of utilizing insects, the amount of pollen is much less; in the highly specialized Orchids, a most successful group, the pollen is reduced to two small bundles.
Once the act of pollination is effected, the duty of the petals and stamens is finished, and they generally fade. The sepals often remain, as in the Rose. By the growth of the pollen tube from the stigma into the ovary, fertilization is effected, and mature seed is produced. The fruit—that is, the seed and its coverings or appendages—offers the most varied forms of any of the plant organs—compare Hazel, Strawberry, Pea, Apple, Cranesbill, Dandelion; the variety is endless. Many of these forms are connected with the means by which seed-dispersal is effected: this subject has been touched on in Chapter III. But in numerous instances we can no more assign a reason for their beautiful or fantastic forms than we can account for the infinite variety of shape assumed by leaves and flowers.
Summing up, then, what has been sketched in this chapter, we must think of our plant as a very complicated and wonderful machine, of which the terrestrial Seed Plant is the highest expression. Water is the basis on which its activities are founded—the currency in which all business is transacted. The amount of water contained in a growing plant is seldom realized. Even solid timber, when growing, is half wood, half water. A fresh lettuce loses 95 per cent. of its weight if the water is driven off by drying. Living in an aerial medium which tends to deprive it of moisture continually, and which furnishes water to the soil only intermittently in the form of rain, and often in sparing quantity, the plant envelops itself from end to end of its exposed portions in a waterproof cuticle; the only openings in its surface layer are the spongy tips of the root hairs on the one hand, and in the stomata on the other. These minutest of openings—so small that the number on a square inch of leaf surface often far exceeds a hundred thousand—might prove danger-points were they not most jealously watched over. But each is provided with a pair of guard-cells ready to close the opening at any moment; and where drought threatens, the whole of the stomata are found in concealed positions. An ample pipe-system extends from root, through stem, to leaf, but it does not communicate directly with the openings at either end. All material, whether liquid or gaseous, absorbed or given out, has to run the gauntlet of the living cells, which are jealous watchmen, and allow only selected substances to pass through them. The crude building materials and food materials are assembled in the leaves, where in cells spread out to the light the chlorophyll is massed. Under the microscope, the chlorophyll is seen to be located in minute granules embedded in the semifluid contents of the cells. Well may we gaze in wonder at these tiny green specks. Each is so small that although a couple of hundred of them are often present in each cell, they occupy but a very small proportion of its volume. The cells themselves are of microscopic size. The chlorophyll itself occupies only quite a small portion of the corpuscle in which it is immersed; yet on its activity as spread in this infinitesimal quantity through the leaves the whole organic world, animal as well as vegetable, depends.[9] Utilizing the energy which comes through space from the sun, it builds up organic compounds; from the energy thus stored comes all the varied life and vital movement which fill our world—the opening of flowers, the hum of insects, the march of armies, and our own restless thought; while its work in the distant past, laid by in coal and oil, warms our houses and drives our trains, factories, and steamships.
The work of the living chlorophyll accomplished, the food materials produced by its agency are sent by the pipe-system to all parts of the plant, for present use, or to be stored in root, stem, or leaf for future requirements.
Nor is our plant the passive, motionless thing that it may appear to be in comparison with animals and their larger movements. Active motion, local and general, though usually of relatively small amount, accompanies all plant-growth. Throughout root, stem, leaf, and flower transference of material is going forward vigorously. The root hairs and stomata are working at high pressure; the chlorophyll never ceases its activities while daylight lasts. Externally, the growing branches, leaves, and flowers also display incessant movement, sweeping the air in small circles, or in the case of climbing plants in curves of considerable amplitude. Alterations of illumination or of temperature produce other movement—bendings towards or away from light, the drooping of leaves and closing of flowers at nightfall, and so on.
All these phenomena of growth and movement culminate in the production of flowers, and in the remarkable developments by which, through the agency of pollen and ovule, a new generation is produced.
The appearance of man upon the Earth is an event of very recent occurrence, not only in terrestrial history, but in the history of organic life in the world. In the life-story which began somewhere in far pre-Cambrian times, the record of the whole of human activities occupies but the last paragraph of the last chapter. For millions of years—ever since the larger animals first abandoned the aquatic haunts of their ancestors and took to a terrestrial life—creatures great and small, of myriad kinds, including huge reptiles and amphibians, and later on a crowd of birds and mammals, have fed on land plants, without effecting any profound changes in the appearance of the mantle of vegetation which covered so much of the Earth’s surface. It has been left for the human race, in the course of the few thousand years that have elapsed since it emerged from an existence comparable to that of the beasts and birds, and learned the arts of peace and war, to effect such sweeping changes in terrestrial vegetation over wide areas, that its influence in this respect requires a separate chapter for its consideration.
The changes referred to are largely—though by no means wholly—due to the requirements of the art of husbandry; and to the history of agriculture we may look for information as to the time and place and nature of man’s conquest of the surface of the globe. At the period of the earliest human civilizations, such as those of Egypt and Mesopotamia, the domestication of plants and animals had already reached an advanced stage. Its origin lies far behind the historic period. We can picture in imagination the time when in all inhabited parts of the globe man wandered with no fixed abode, seeking food when he was hungry, and making no provision for the morrow. Residence in a spot which afforded a valued supply of food, such as an abundance of buckwheat or millet or dates or bread-fruit, might lead to a desire to encourage the growth of such useful plants by protecting them and their offspring; following on which might arise the practice of assisting their growth, and thus eventually of cultivating them. Selection of the most productive strains would gradually follow, and barter would cause the spread of useful plants over wider and wider areas. We can picture development from such rude beginnings into the regular cultivation of the soil and the enclosing of the cultivated areas for their protection. It is clear that such practices would not readily arise among nomadic tribes, nor among those inhabiting forest regions where the ground was densely covered by trees. An abundance of animal food would produce a race of hunters rather than of tillers of the soil; and as for forest regions, they are unsuitable for human development; forest races have never been pioneers of civilization. Before agriculture—indeed, before civilization in any form—could make much progress, a settled life was necessary, free from migrations in search of food or for the avoidance of enemies. Hence the earliest civilizations tended to arise in areas which were protected by natural ramparts from the irruption of rival tribes. Egypt had the desert on three sides, and the sea—an impassable barrier to early peoples—on the fourth. The valleys of the Euphrates and Tigris presented similar features. In both areas rich alluvial soil offered a full reward to attempts at agriculture, and the alternation of summer and winter encouraged the making of provision for the non-productive period by the taking advantage of the period of growth: conditions not present under the “endless summer skies” of Tropical lands, where an easy and perennial food-supply tended against the development of industry.
The basin of the Mediterranean—the cradle of the earlier Western civilizations from the time of Egypt down to Rome—was, then, also the cradle of European agriculture. These lands, with their wet winters and dry summers, the latter inimical to the development of tree growth, lent themselves to cultivation more readily than the great forest-belt which lay to the northward, sweeping across Europe from Britain to the Urals. Although there is clear evidence that grain was cultivated in Europe as far back as the Neolithic Period (say 7,000 to 5,000 B.C.), it seems established that when Roman agriculture stood at its perfection the peoples to the north were still mainly nomads, dependent for their food-supply on their flocks or on the chase. In Britain, Cæsar found corn grown in Southern England, but the centre and north were largely forest land tenanted by tribes living on flesh and milk, and clothed in skins. The vigorous colonization of the Romans may well have been accountable for the introduction into Britain of many of the farm plants still grown there. The wars of the next fifteen hundred years on the one hand, and the spread of agriculture on the other, caused the steady destruction of the forests, till at length England and Central Europe began to assume their present appearance. The draining of marshes and fens, the enclosing of land, went on steadily, and to a slight extent is going on still; within recent years, the European War has resulted in the disappearance of many of the remaining woods, and in the breaking up of fresh land.
From the point of view of the botanist, agriculture consists of the destruction of the plant associations which for some thousands of years have occupied the ground, and their replacement by other plants which are useful to man. The natural plant associations being the result of the survival of the fittest through a long period of time, while the farmer’s crops represent plants which do not grow naturally on the ground, nor often indeed in the country (while they are frequently artificial forms unable to reproduce themselves), it follows that the latter cannot compete with the former, and can be maintained only by the most careful protection. The native plants are always striving to reoccupy their legitimate territory, and the farmer is incessantly engaged in trying to keep them out. Agriculture, indeed, has been defined as “a controversy with weeds.” Incidentally, the suppression of the natural flora allows many weaker plants an opportunity of which they are not slow to take advantage. These may be natives, but are often annuals which have followed the spread of farming operations, or which are directly—though unintentionally—introduced by man as impurities in the seed which he sows.
Let us look a little more closely into the question of profit and loss in our flora resulting from agriculture. In the first place, whether the ground is tilled or grazed, the woodland which primitively occupied so much of it disappears. The plough and the scythe are fatal to all seedling trees. Little less fatal is the browsing of cattle and sheep, and even in rough pasture only thorny plants like Whitethorn and Gorse may be found battling successfully for a lodgment. Where woodland is used for pasturage, the delicate shade plants—Anemones, Wild Hyacinths, Primroses—soon die out. No young trees appear on the grazed surface, though hundreds of thousands of seeds may be shed annually over the ground. In the course of time the present trees will die, and only grass remain. How different is it where cattle are excluded and the scythe unused! Among the grass young trees spring up everywhere, and in the woods a dense undergrowth of saplings sheltering a varied shade flora makes its appearance; regeneration of the natural woodland proceeds apace.
Natural grasslands, if undisturbed, possess a flora which has been built up during a long period of time, and which, like all purely natural plant associations, represents a delicate balance between its many constituents, which often include rare and shy species. If such land be once broken up, its flora will probably never again resume its former composition even if allowed to regenerate during a long series of years, for the alteration in the old substratum caused by its being turned over and mixed introduces new edaphic (i.e., soil) conditions which will not entirely pass away. As regards grazing, likewise, when land is pastured up to or near its full capacity, as is generally the case on enclosed areas, the weaker and often more interesting members of the flora tend to disappear. In primitive times all grasslands had, of course, their natural grazing inhabitants—in our islands deer of more than one species, sheep, and smaller creatures such as rabbits and geese—and so a total exclusion of grazing animals now would no more tend to reproduce exactly the flora of pre-husbandry days than does the excess of herbivores; but the present heavy stocking of the land is to be deplored by the botanist, even as it is rejoiced in by the economist. The more vigorous plants, and especially those which propagate themselves largely by vegetative means, survive, or even increase owing to the augmented food supplied by the manure which the animals provide; but many species fail to ripen seed, being either eaten or trampled; the rarer Orchids, strange ferns like the Adder’s Tongue (Ophioglossum vulgatum), and Moonwort (Botrychium Lunaria), and the other choicer denizens of the grasslands, tend to disappear.
Drainage is an obvious cause of loss to our flora. Whole lakes and areas of swamp, with their peculiar and to a great extent natural flora, have disappeared from parts of the country. Some of the most interesting marsh plants of the British flora—such as the two fine Ragweeds, Senecio palustris and S. paludosus, and the Marsh Sow-thistle (Sonchus palustris)—have on this account almost vanished from our islands, like the Bittern and Great Bustard which are their companions.
Some lakes, again, have been ruined for the botanist by being used as reservoirs. The considerable changes of level which this involves is a thing to which plants are not adapted, and only a few can withstand it, such as the Water Bistort (Polygonum amphibium) and the Shore-weed (Littorella uniflora), which are equally at home on land or in water, being able to change rapidly their structure and mode of life to suit change of environment. As compared with a lakelet with a natural outlet, a dam with a sluice has always a much reduced and usually quite uninteresting flora.
The proximity of a large town, especially if it is a centre of manufacture, is a notorious factor in the reduction of the native flora: not only by the thoughtless and wanton destruction carried out by its inhabitants, but more subtly by the deposition of soot, and by the poisoning of the air by sulphurous and acid fumes. The higher Cryptogams, such as Mosses and Hepatics, are particularly susceptible in this respect, and vanish along with the more delicate Seed Plants. Mining centres are specially destructive of plant life, since, in addition to other drawbacks, the soil is often buried under masses of excavated material containing poisonous substances. If there is a purgatory for plants, it is surely found in such areas.
Other examples of the multitudinous ways in which human activities disturb and destroy native plant life will occur to the reader—the burning of moors in order to improve them as pasturage; in recent years the tarring of roads, which kills the pleasant wayside herbage and poisons the streams into which the road drainage is carried; and so on. The indictment is an overwhelming one, and, as said in the first chapter, the flora is now everywhere so altered that we can gain some idea of its original aspect only by a study of isolated fragments and much-adulterated samples.
But if the debit side of the account, as presented by the lover of nature, is heavy, it must not be forgotten that there are many items to man’s credit. Though our country’s vegetation has lost in scientific interest, it has gained vastly in both economic and æsthetic value by the introduction of useful and ornamental plants from all the Temperate regions of the world; and besides, a large number of species have followed in man’s footsteps, and, taking advantage of the disturbance of the native flora caused by his operations, endeavour with more or less success to establish a footing in the country. Before we trespass on the domains of arboriculture, horticulture, or agriculture, under which heads the cultivation of useful or ornamental plants divides itself, some consideration is required of those plants which, quasi-wild, are usually included in accounts of the vegetation under the head of aliens, denizens, colonists, and so forth. These constitute a quite considerable proportion of the total number of species found in any area which has felt the influence of man. For instance, in the county of Dublin, which, owing to its diversified surface—sea-cliff, sands, moorland, woodland, and cultivation—and its favourable climate—the warmest and driest in the country—possesses the largest flora of any similar area (354 square miles) in Ireland, the list of about 760 “wild” plants includes some 170, or over one-fifth of the whole, whose presence is attributable, directly or indirectly, to human activities. We may compare these figures with those drawn from a study of the flora of Kent, which faces across the Channel towards France just as county Dublin faces across the Irish Sea towards England; both are areas of early settlement and both lie in the main stream of traffic. In Kent we have to deal with a larger area (1,570 square miles), and a larger flora (1,160 species). We find that, of these 1,160 species, 146, or about one-eighth, are set down as owing their presence to man.[10] And so it is in all the more populous and highly tilled parts of our islands.
This question of alien plants, their past history and present standing, is one of the most puzzling with which the student of our flora has to deal. In the first place, most of them have been in the country for a long time, and the record of their introduction is lost. Next, while many of them are confined to ground disturbed by man, and thus clearly exist under man’s protection—however unwillingly that protection may be afforded—others have mixed with the indigenous flora, won a place in the closed native vegetation, and might be ranked as true natives were it not that a study of their general distribution raises doubts as to the possibility of their having arrived in our islands unaided—doubts which their known occurrence in gardens tends to confirm. Take the case of the Yellow Monkey-flower (Mimulus Langsdorfii). This has quite established itself in our native flora, in some places ascending mountain streams far into the hills, in others mingling with the rank flora of muddy estuaries. It looks as aboriginal as any of the plants among which it grows: but the facts that the genus to which it belongs is American (with a few species in Australia and New Zealand), that it itself is found native in the western States and not in the eastern, and that it has been long cultivated in gardens, furnish convincing proof that it is really an alien. But it is seldom that the evidence is so satisfactory as in this case. More usually the range of the doubtful members of the flora is continuous, extending from regions where they are truly native to others where they are undoubtedly exotic. For instance, many annual plants of the Mediterranean region have followed the spread of agriculture across the former forest areas of Central and Western Europe into our own islands. Plants native in France have been transported into England, and English natives into Ireland; east Irish plants have spread westward—sixty years ago, save for a single record of P. hybridum, Papaver dubium was the only Poppy known west of the Shannon; now all four British species occur, several of them in many places. The flora of Europe, as pointed out already, diminishes in variety as we pass westward into the outlying areas. Those species whose aboriginal distribution stopped short of the western limit of the land had no doubt a fluctuating western or northern or southern boundary to their range, dependent on temporary conditions. Thus, a hard winter might kill back a plant already at the limit of its natural range, or a warm summer, by ripening abundance of its seed, might result in its slight advance. The general effect of human operations has been to lessen competition and increase suitable habitats by the destruction of the native vegetation which occupied them, and this has resulted in a general advance of a large number of species. What renders the study of this advance so difficult is the fact that on all disturbed land the truly native plants which have been ousted are striving side by side with the immigrants to regain their former territories; and it is now often very difficult to disentangle them: to separate the sheep from the goats. If only we could have had a Watson’s “Topographical Botany” written five thousand years ago, before our restless race began to mess up the vegetation!
However, as has been said, what we have lost on one side we have gained on another. On every side bright immigrants meet the eye. Our old buildings and quarries often blaze with the Red Valerian (Kentranthus ruber) and Wallflower (Cheiranthus Cheiri); in fields Poppies of various kinds, Corn Cockle (Lychnis Githago), and Corn Blue-bottle (Centaurea Cyanus) add a glory to the rich green or gold of the cereals; dry banks and gravelly places are decorated with species of Melilot (Melilotus), Chamomile (Anthemis), Knapweed (Centaurea), and many others. The flora of harbours and docksides is often as cosmopolitan as the sailors of the ships by whose agency it came there; and the unfamiliar weeds—the gipsies and tramps of the plant world—which we encounter on roadsides, rubbish-heaps, and railway stations lend an additional interest to our botanical rambles.
Turning now to the plants which are used by man, it may be pointed out in the first place that the human race obtains much more, whether of profit or of pleasure, from the vegetable than from the animal kingdom. Flesh, whether derived from mammals, birds, or fishes; wool, silk, leather, oils, and so on, bulk much less than the grains, vegetables, fruits, timber, fibres, fodder plants, and other vegetable products which we use in our daily life. On the æsthetic side, again, while the beauty of birds and insects is a source of frequent delight, flowers play a part in daily life that the more delicate and sensitive animals can never do. Again, in the number of different species used, whether for profit or pleasure, the plant world takes precedence. This is especially the case as regards our farms and pleasure grounds, plants lending themselves much more readily to domestication than animals do. And so a suburban house may have a hundred or a thousand different plants in kitchen garden and flower plot, orchard, and shrubbery, while its animal dependents consist of a horse, a couple of dogs, a cat, some fowl, and a canary. So again a Botanic Garden may easily possess as many thousands of different species as a Zoological Garden contains hundreds.
This army of plants which human beings collect about themselves may be grouped under two categories—useful and ornamental. On a previous page (p. 136) a suggestion has been made as to how the cultivation of useful plants may have arisen. As now practised, this industry is the largest in the world, and with the growth of means of transport has ceased to be only or even mainly of local importance: we use every day wheat from Australia, rice from China, tea from India, cotton from the United States, timber from Norway. In some cases, as in the last, these materials are harvested as they occur in the wild state, but in the majority of instances the plants are not merely conserved, but cultivated; cultivation has led to selection of the best varieties; and continued selection has resulted in the production of forms often very different in appearance from the wild plants from which they originated. We cannot create new forms; but by taking advantage of the innate tendency to vary which all plants display—some to a much greater degree than others—and by raising, generation after generation, the seeds of those individuals in which a certain abnormal feature is best displayed, we can produce an artificial race in which the selected character may be developed to an extraordinary degree. But we have not by this means produced a new species. Seedlings of such plants will tend to “throw back” towards the original form; we can preserve or improve the special characters only by continued selection; if allowed to grow and seed unchecked, most of such plants will revert to the natural type in a few generations. Often this reversion is so rapid that seeds are useless for cultural purposes, and it is only by cuttings or graftings—that is, by growing parts of the original possessor of the required characters—that constancy can be maintained; this is what is usually done in the case of fruit-trees, Roses, Pansies, and so on.
Equally efficient in the hands of the cultivator has been another method of producing new forms—namely, hybridization. If the pollen of a plant be transferred to the stigma of a related species, offspring is often produced; and the product is a batch of plants intermediate in characters between the two parents, and generally uniform in appearance. Should these be crossed again, a heterogeneous offspring is the result, displaying a variety of characters inherited from one or other original parent. The crossing of varieties, native or cultivated, has the same result. Hybrids occur in nature, but not very frequently. Insects visiting flowers are well known to confine their attention to a great extent to one species at a time, so as agents of hybridization they are not efficient. Again, many hybrids do not produce fertile seed, so that if they arise by natural means they are not perpetuated. In the garden, hybridizing has been resorted to largely; but its practice is not so ancient as the method of producing improved breeds by selection.
The cultivation of specially selected forms is certainly of remote origin, and probably goes back to the earliest days of agriculture: of early date, too, is the introduction into regions where they do not occur naturally of plants desirable for their use or beauty. The records of the cultivation of the Vine, for instance, go back for five or six thousand years in Egypt. Two thousand years ago Pliny writes that ninety-one principal forms could be reckoned in his day, though “the varieties are very nearly as numberless as the districts in which they grow.” Theophrastus, three hundred years earlier, discourses learnedly of the different kinds of cultivated Figs, etc., and their superiority over the wild kinds. These and other authors make frequent mention of plants introduced into Greece or Italy from the East for their usefulness or their pleasing qualities. Nowadays, the number of species cultivated, the innumerable forms of these which are grown, and the wide distribution which these forms have attained, have resulted in the cultivated flora of a country like England being, so far as the higher plants are concerned, much larger than the native flora, even when all the plants which are grown under glass are left out of consideration.
In the case of plants of economic importance, the usual aim of selection has been increase of size or productiveness of the parts which are useful. In some instances selection has taken several directions inside the limits of a single species, as in the forms of Cabbage, which are all the offspring of Brassica oleracea (Fig. 25), a seaside plant of Western and Southern Europe, and are mostly creations of comparatively recent date. The Cauliflower has been produced by increasing the size of the inflorescence; White Cabbage by promoting leaf production; Brussels Sprouts by encouraging the development of axillary shoots; while a form with a tall and woody stem is made into walking sticks. More often we find a species developed along a single line. For instance, the tendency to store food materials in a fleshy taproot has been developed in the case of Turnip, Beet, Carrot; the fleshy scale-leaves which form bulbs have been exploited in the case of the Onion; increased stem-growth is promoted in Asparagus; increased leaf-growth in Spinach and Lettuce; while by the development of seeds and fruits of many kinds artificial selection has supplied us with the foods on which the human race mainly subsists. The most important of all these last are, of course, the different grains, which are the seeds of grasses of various genera—Triticum (Wheat), Hordeum (Barley), Secale (Rye), Avena (Oat), Panicum (Millet), Oryza (Rice),