Botany

Botany (from Gr. βοτάνη, plant; βόσκειν, to graze), the science which includes everything relating to the vegetable kingdom, whether in a living or in a fossil state. It embraces a consideration of the external forms of plants—of their anatomical structure, however minute—of the functions which they perform—of their arrangement and classification—of their distribution over the globe at the present and at former epochs—and of the uses to which they are subservient. It examines the plant in its earliest state of development, and follows it through all its stages of progress until it attains maturity. It takes a comprehensive view of all the plants which cover the earth, from the minutest organism, only visible by the aid of the microscope, to the most gigantic productions of the tropics. It marks the relations which subsist between all members of the plant world, including those between existing groups and those which are known only from their fossilized remains preserved in the rocks. We deal here with the history and evolution of the science.

The plants which adorn the globe more or less in all countries must necessarily have attracted the attention of mankind from the earliest times. The science that treats of them dates back to the days of Solomon, who “spake of trees, from the cedar of Lebanon to the hyssop on the wall.” The Chaldaeans, Egyptians and Greeks were the early cultivators of science, and botany was not neglected, although the study of it was mixed up with crude speculations as to vegetable life, and as to the change of plants into animals. About 300 years before Christ Theophrastus wrote a History of Plants, and described about 500 species used for the treatment of diseases. Dioscorides, a Greek writer, who appears to have flourished about the time of Nero, issued a work on Materia Medica. The elder Pliny described about a thousand plants, many of them famous for their medicinal virtues. Asiatic and Arabian writers also took up this subject. Little, however, was done in the science of botany, properly so called, until the 16th century of the Christian era, when the revival of learning dispelled the darkness which had long hung over Europe. Otto Brunfels, a physician of Bern, has been looked upon as the restorer of the science in Europe. In his Herbarium, printed at Strassburg (1530-1536), he gave descriptions of a large number of plants, chiefly those of central Europe, illustrated by beautiful woodcuts. He was followed by other writers,—Leonhard Fuchs, whose Historia Stirpium (Basel, 1542) is worthy of special note for its excellent woodcuts; Hieronymus Bock, whose Kreutter Buch appeared in 1539; and William Turner, “The Father of English Botany,” the first part of whose New Herbal, printed in English, was issued in 1551. The descriptions in these early works were encumbered with much medicinal detail, including speculations as to the virtues of plants. Plants which were strikingly alike were placed together, but there was at first little attempt at systematic classification. A crude system, based on the external appearance of plants and their uses to man, was gradually evolved, and is well illustrated in the Herbal, issued in 1597 by John Gerard (1545-1612), a barber-surgeon, who had a garden in Holborn, and was a keen student of British plants.

One of the earliest attempts at a methodical arrangement of plants was made in Florence by Andreas Caesalpinus (1519-1603), who is called by Linnaeus primus verus systematicus. In his work De Plantis, published at Florence in 1583, he distributed the 1520 plants then known into fifteen classes, the distinguishing characters being taken from the fruit.

John Ray (1627-1705) did much to advance the science of botany, and was also a good zoologist. He promulgated a system which may be considered as the dawn of the “natural system” of the present day (Ray, Methodus Plantarum, 1682). He separated flowering from flowerless plants, and divided the former into Dicotyledons and Monocotyledons. His orders (or “classes”) were founded to some extent on a correct idea of the affinities of plants, and he far outstripped his contemporaries in his enlightened views of arrangement.

About the year 1670 Dr Robert Morison1 (1620-1683), the first professor of botany at Oxford, published a systematic arrangement of plants, largely on the lines previously suggested by Caesalpinus. He divided them into eighteen classes, distinguishing plants according as they were woody or herbaceous, and taking into account the nature of the flowers and fruit. In 1690 Rivinus2 promulgated a classification founded chiefly on the forms of the flowers. J.P. de Tournefort3 (1656-1708), who about the same time took up the subject of vegetable taxonomy, was long at the head of the French school of botany, and published a systematic arrangement in 1694-1700. He described about 8000 species of plants, and distributed them into twenty-two classes, chiefly according to the form of the corolla, distinguishing herbs and under-shrubs on the one hand from trees and shrubs on the other. The system of Tournefort was for a long time adopted on the continent, but was ultimately displaced by that of Carl von Linné, or Linnaeus (q.v.; 1707-1778).

The system of Linnaeus was founded on characters derived from the stamens and pistils, the so-called sexual organs of the flower, and hence it is often called the sexual system. It is an artificial method, because it takes into account only a few marked characters in plants, and does not propose to unite them by natural affinities. It is an index to a department of the book of nature, and as such is useful to the student. It does not aspire to any higher character, and although it cannot be looked upon as a scientific and natural arrangement, still it has a certain facility of application which at once commended it. It does not of itself give the student a view of the true relations of plants, and by leading to the discovery of the name of a plant, it is only a stepping-stone to the natural system. Linnaeus himself claimed nothing higher for it. He says—“Methodi Naturalis fragmenta studiose inquirenda sunt. Primum et ultimum hoc in botanicis desideratum est. Natura non facit saltus. Plantae omnes utrinque affinitatem monstrant, uti territorium in mappa geographica.” Accordingly, besides his artificial index, he also promulgated fragments of a natural method of arrangement.

The Linnean system was strongly supported by Sir James Edward Smith (1759-1828), who adopted it in his English Flora, and who also became possessor of the Linnean collection. The system was for a long time the only one taught in the schools of Britain, even after it had been discarded by those in France and in other continental countries.

The foundation of botanic gardens during the 16th and 17th centuries did much in the way of advancing botany. They were at first appropriated chiefly to the cultivation of medicinal plants. This was especially the case at universities, where medical schools existed. The first botanic garden was established at Padua in 1545, and was followed by that of Pisa. The garden at Leiden dates from 1577, that at Leipzig from 1579. Gardens also early existed at Florence and Bologna. The Montpellier garden was founded in 1592, that of Giessen in 1605, of Strassburg in 1620, of Altdorf in 1625, and of Jena in 1629. The Jardin des Plantes at Paris was established in 1626, and the Upsala garden in 1627. The botanic garden at Oxford was founded in 1632\. The garden at Edinburgh was founded by Sir Andrew Balfour and Sir Robert Sibbald in 1670, and, under the name of the Physic Garden, was placed under the superintendence of James Sutherland, afterwards professor of botany in the university. The garden at Kew dates from about 1730, when Frederick, prince of Wales, obtained a long lease of Kew House and its gardens from the Capel family. After his death in 1751 his widow, Princess Augusta of Saxe-Gotha, showed great interest in their scientific development, and in 1759 engaged William Aiton to establish a Physic Garden. The garden of the Royal Dublin Society at Glasnevin was opened about 1796; that of Trinity College, Dublin, in 1807; and that of Glasgow in 1818. The Madrid garden dates from 1763, and that of Coimbra from 1773. Jean Gesner (1709-1790), a Swiss physician and botanist, states that at the end of the 18th century there were 1600 botanic gardens in Europe.

A new era dawned on botanical classification with the work of Antoine Laurent de Jussieu (1748-1836). His uncle, Bernard de Jussieu, had adopted the principles of Linnaeus’s Fragmenta in his arrangement of the plants in the royal garden at the Trianon. At an early age Antoine became botanical demonstrator in the Jardin des Plantes, and was thus led to devote his time to the science of botany. Being called upon to arrange the plants in the garden, he necessarily had to consider the best method of doing so, and, following the lines already suggested by his uncle, adopted a system founded in a certain degree on that of Ray, in which he embraced all the discoveries in organography, adopted the simplicity of the Linnean definitions, and displayed the natural affinities of plants. His Genera Plantarum, begun in 1778, and finally published in 1789, was an important advance, and formed the basis of all natural classifications. One of the early supporters of this natural method was Augustin Pyramus de Candolle (1778-1841), who in 1813 published his Théorie élémentaire de la botanique, in which he showed that the affinities of plants are to be sought by the comparative study of the form and development of organs (morphology), not of their functions (physiology). His Prodromus Systematis Naturalis Regni Vegetabilis was intended to embrace an arrangement and description of all known plants. The work was continued after his death, by his son Alphonse de Candolle, with the aid of other eminent botanists, and embraces descriptions of the genera and species of the orders of Dicotyledonous plants. The system followed by de Candolle is a modification of that of Jussieu.

In arranging plants according to a natural method, we require to have a thorough knowledge of structural and morphological botany, and hence we find that the advances made in these departments have materially aided the efforts of systematic botanists.

Robert Brown (1773-1858) was the first British botanist to support and advocate the natural system of classification. The publication of his Prodromus Florae Novae Hollandiae (in 1810), according to the natural method, led the way to the adoption of that method in the universities and schools of Britain. In 1827 Brown announced his important discovery of the distinction between Angiosperms and Gymnosperms, and the philosophical character of his work led A. von Humboldt to refer to him as “Botanicorum facile princeps.” In 1830 John Lindley published the first edition of his Introduction to the Natural System, embodying a slight modification of de Candolle’s system. From the year 1832 up to 1859 great advances were made in systematic botany, both in Britain and on the continent of Europe. The Enchiridion and Genera Plantarum of S.L. Endlicher (1804-1849), the Prodromus of de Candolle, and the Vegetable Kingdom (1846) of J. Lindley became the guides in systematic botany, according to the natural system.

The least satisfactory part of all these systems was that concerned with the lower plants or Cryptogams as contrasted with the higher or flowering plants (Phanerogams). The development of the compound microscope rendered possible the accurate study of their life-histories; and the publication in 1851 of the results of Wilhelm Hofmeister’s researches on the comparative embryology of the higher Cryptogamia shed a flood of light on their relationships to each other and to the higher plants, and supplied the basis for the distinction of the great groups Thallophyta, Bryophyta, Pteridophyta and Phanerogamae, the last named including Gymnospermae and Angiospermae.

A system of classification for the Phanerogams, or, as they are frequently now called, Spermatophyta (seed-plants), which has been much used in Great Britain and in America, is that of Bentham and Hooker, whose Genera Plantarum (1862-1883) is a descriptive account of all the genera of flowering plants, based on their careful examination. The arrangement is a modification of that adopted by the de Candolles. Another system differing somewhat in detail is that of A.W. Eichler (Berlin, 1883), a modified form of which was elaborated by Dr Adolf Engler of Berlin, the principal editor of Die natrürliche Pflanzenfamilien.

The study of the anatomy and physiology of plants did not keep pace with the advance in classification. Nehemiah Grew and his contemporary Marcello Malpighi were the earliest discoverers in the department of plant anatomy. Both authors laid an account of the results of their study of plant structure before the Royal Society of London almost at the same time in 1671. Malpighi’s complete work, Anatome Plantarum, appeared in 1675 and Grew’s Anatomy of Plants in 1682. For more than a hundred years the study of internal structure was neglected. In 1802 appeared the Traité d’anatomie et de physiologie végétale of C.F.B. de Mirbel (1776-1854), which was quickly followed by other publications by Kurt Sprengel, L.C. Treviranus (1779-1864), and others. In 1812 J.J. P. Moldenhawer isolated cells by maceration of tissues in water. The work of F.J.F. Meyen and H. von Mohl in the middle of the 19th century placed the study of plant anatomy on a more scientific basis. Reference must also be made to M.J. Schleiden (1804-1881) and F. Unger (1800-1870), while in K.W. von Nägeli’s investigations on molecular structure and the growth of the cell membrane we recognize the origin of modern methods of the study of cell-structure included under cytology (q.v.). The work of Karl Sanio and Th. Hartig advanced knowledge on the structure and development of tissues, while A. de Bary’s Comparative Anatomy of the Phanerogams and Ferns (1877) supplied an admirable presentation of the facts so far known. Since then the work has been carried on by Ph. van Tieghem and his pupils, and others, who have sought to correlate the large mass of facts and to find some general underlying principles (see Plants: Anatomy of).

The subject of fertilization was one which early excited attention. The idea of the existence of separate sexes in plants was entertained in early times, long before separate male and female organs had been demonstrated. The production of dates in Egypt, by bringing two kinds of flowers into contact, proves that in very remote periods some notions were entertained on the subject. Female date-palms only were cultivated, and wild ones were brought from the desert in order to fertilize them. Herodotus informs us that the Babylonians knew of old that there were male and female date-trees, and that the female required the concurrence of the male to become fertile. This fact was also known to the Egyptians, the Phoenicians and other nations of Asia and Africa. The Babylonians suspended male clusters from wild dates over the females; but they seem to have supposed that the fertility thus produced depended on the presence of small flies among the wild flowers, which, by entering the female flowers, caused them to set and ripen. The process was called palmification. Theophrastus, who succeeded Aristotle in his school in the 114th Olympiad, frequently mentions the sexes of plants, but he does not appear to have determined the organs of reproduction. Pliny, who flourished under Vespasian, speaks particularly of a male and female palm, but his statements were not founded on any real knowledge of the organs. From Theophrastus down to Caesalpinus, who died at Rome in 1603, there does not appear to have been any attention paid to the reproductive organs of plants. Caesalpinus had his attention directed to the subject, and he speaks of a halitus or emanation from the male plants causing fertility in the female.

Nehemiah Grew seems to have been the first to describe, in a paper on the Anatomy of Plants, read before the Royal Society in November 1676, the functions of the stamens and pistils. Up to this period all was vague conjecture. Grew speaks of the attire, or the stamens, as being the male parts, and refers to conversations with Sir Thomas Millington, Sedleian professor at Oxford, to whom the credit of the sexual theory seems really to belong. Grew says that “when the attire or apices break or open, the globules or dust falls down on the seedcase or uterus, and touches it with a prolific virtue.” Ray adopted Grew’s views, and states various arguments to prove their correctness in the preface to his work on European plants, published in 1694. In 1694 R.J. Camerarius, professor of botany and medicine at Tübingen, published a letter on the sexes of plants, in which he refers to the stamens and pistils as the organs of reproduction, and states the difficulties he had encountered in determining the organs of Cryptogamic plants. In 1703 Samuel Morland, in a paper read before the Royal Society, stated that the farina (pollen) is a congeries of seminal plants, one of which must be conveyed into every ovum or seed before it can become prolific. In this remarkable statement he seems to anticipate in part the discoveries afterwards made as to pollen tubes, and more particularly the peculiar views promulgated by Schleiden. In 1711 E.F. Geoffrey, in a memoir presented to the Royal Academy at Paris, supported the views of Grew and others as to the sexes of plants. He states that the germ is never to be seen in the seed till the apices (anthers) shed their dust; and that if the stamina be cut out before the apices open, the seed will either not ripen, or be barren if it ripens. He mentions two experiments made by him to prove this—one by cutting off the staminal flowers in Maize, and the other by rearing the female plant of Mercurialis apart from the male. In these instances most of the flowers were abortive, but a few were fertile, which he attributes to the dust of the apices having been wafted by the wind from other plants.

Linnaeus took up the subject in the inauguration of his sexual system. He first published his views in 1736, and he thus writes—“Antheras et stigmata constituere sexum plantarum, a palmicolis, Millingtono, Grewio, Rayo, Camerario, Godofredo, Morlando, Vaillantio, Blairio, Jussievio, Bradleyo, Royeno, Logano, &c., detectum, descriptum, et pro infallibili assumptum; nec ullum, apertis oculis considerantem cujuscunque plantae flores, latere potest.” He divided plants into sexual and asexual, the former being Phanerogamous or flowering, and the latter Cryptogamous or flowerless. In the latter division of plants he could not detect stamens and pistils, and he did not investigate the mode in which their germs were produced. He was no physiologist, and did not promulgate any views as to the embryogenic process. His followers were chiefly engaged in the arrangement and classification of plants, and while descriptive botany made great advances the physiological department of the science was neglected. His views were not, however, adopted at once by all, for we find Charles Alston stating arguments against them in his Dissertation on the Sexes of Plants. Alston’s observations were founded on what occurred in certain unisexual plants, such as Mercurialis, Spinach, Hemp, Hop and Bryony. The conclusion at which he arrives is that the pollen is not in all flowering plants necessary for impregnation, for fertile seeds can be produced without its influence. He supports parthenogenesis in some plants. Soon after the promulgation of Linnaeus’s method of classification, the attention of botanists was directed to the study of Cryptogamic plants, and the valuable work of Johann Hedwig (1730-1799) on the reproductive organs of mosses made its appearance in 1782. He was one of the first to point out the existence of certain cellular bodies in these plants which appeared to perform the functions of reproductive organs, and to them the names of antheridia and pistillidia were given. This opened up a new field of research, and led the way in the study of Cryptogamic reproduction, which has since been much advanced by the labours of numerous botanical inquiries. The interesting observations of Morland, already quoted, seem to have been neglected, and no one attempted to follow in the path which he had pointed out. Botanists were for a long time content to know that the scattering of the pollen from the anther, and its application to the stigma, were necessary for the production of perfect seed, but the stages of the process of fertilization remained unexplored. The matter seemed involved in mystery, and no one attempted to raise the veil which hung over the subject of embryogeny. The general view was, that the embryo originated in the ovule, which was in some obscure manner fertilized by the pollen.

In 1815 L.C. Treviranus, professor of botany in Bonn, roused the attention of botanists to the development of the embryo, but although he made valuable researches, he did not add much in the way of new information. In 1823 G.B. Amici discovered the existence of pollen tubes, and he was followed by A.T. Brongniart and R. Brown. The latter traced the tubes as far as the nucleus of the ovule. These important discoveries mark a new epoch in embryology, and may be said to be the foundation of the views now entertained, which were materially aided by the subsequent elucidation of the process of cytogenesis, or cell-development, by Schleiden, Schwann, Mohl and others. The whole subject of fertilization and development of the embryo has been more recently investigated with great assiduity and zeal, as regards both cryptogamous and phanerogamous plants, and details must be sought in the various special articles. The observations of Darwin as to the fertilization of orchids, Primula, Linum and Lythrum, and other plants, and the part which insects take in this function, gave an explanation of the observations of Christian Konrad Sprengel, made at the close of the 18th century, and opened up a new phase in the study of botany, which has been followed by Hermann Müller, Federico Delpino and others, and more recently by Paul Knuth.

One of the earliest workers at plant physiology was Stephen Hales. In his Statical Essays (1727) he gave an account of numerous experiments and observations which he had made on the nutrition of plants and the movement of sap in them. He showed that the gaseous constituents of the air contribute largely to the nourishment of plants, and that the leaves are the organs which elaborate the food; the importance of leaves in nutrition had been previously pointed out by Malpighi in a short account of nutrition which forms an appendix to his anatomical work. The birth of modern chemistry in the work of J. Priestley and Lavoisier, at the close of the 18th century, made possible the scientific study of plant-nutrition, though Jan Ingenhousz in 1779 discovered that plants incessantly give out carbonic acid gas, but that the green leaves and shoots only exhale oxygen in sunlight or clear daylight, thereby indicating the distinction between assimilation of carbonic acid gas (photosynthesis) and respiration. N.T. de Saussure (1767-1845) gave precision to the science of plant-nutrition by use of quantitative methods. The subjects of plant nutrition and respiration were further studied by R.J.H. Dutrochet towards the middle of the century, and Liebig’s application of chemistry to agriculture and physiology put beyond question the parts played by the atmosphere and the soil in the nutrition of plants.

The phenomena of movements of the organs of plants attracted the attention of John Ray (1693), who ascribed the movements of the leaf of Mimosa and others to alteration in temperature. Linnaeus also studied the periodical movements of flowers and leaves, and referred to the assumption of the night-position as the sleep-movement. Early in the 19th century Andrew Knight showed by experiment that the vertical growth of stems and roots is due to the influence of gravitation, and made other observations on the relation between the position assumed by plant organs and external directive forces, and later Dutrochet, H. von Mohl and others contributed to the advance of this phase of plant physiology. Darwin’s experiments in reference to the movements of climbing and twining plants, and of leaves in insectivorous plants, have opened up a wide field of inquiry as to the relation between plants and the various external factors, which has attracted numerous workers. By the work of Julius Sachs and his pupils plant physiology was established on a scientific basis, and became an important part of the study of plants, for the development of which reference may be made to the article Plants: Physiology. The study of form and development has advanced under the name “morphology,” with the progress of which are associated the names of K. Goebel, E. Strasburger, A. de Bary and others, while more recently, as cytology (q.v.), the intimate study of the cell and its contents has attracted considerable attention.

The department of geographical botany made rapid advance by means of the various scientific expeditions which have been sent to all quarters of the globe, as well as by individual effort (see Plants: Distribution) since the time of A. von Humboldt. The question of the mode in which the floras of islands and of continents have been formed gave rise to important speculations by such eminent botanical travellers as Charles Darwin, Sir J.D. Hooker, A.R. Wallace and others. The connexion between climate and vegetation has also been studied. Quite recently under the name of “Ecology” or “Oecology” the study of plants in relation to each other and to their environment has become the subject of systematic investigation.

The subject of palaeontological botany (see Palaeobotany) has been advanced by the researches of both botanists and geologists. The nature of the climate at different epochs of the earth’s history has also been determined from the character of the flora. The works of A.T. Brongniart, H.R. Goeppert and W.P. Schimper advanced this department of science. Among others who contributed valuable papers on the subject may be noticed Oswald Heer (1809-1883), who made observations on the Miocene flora, especially in Arctic regions; Gaston de Saporta (1823-1895), who examined the Tertiary flora; Sir J.W. Dawson and Leo Lesquereux, and others who reported on the Canadian and American fossil plants. In Great Britain also W.C. Williamson, by his study of the structure of the plants of the coal-measures, opened up a new line of research which has been followed by Bertrand Renault, D.H. Scott, A.C. Seward and others, and has led to important discoveries on the nature of extinct groups of plants and also on the phylogeny of existing groups.

Botany may be divided into the following departments:—

1\. Structural, having reference to the form and structure of the various parts, including (a) Morphology, the study of the general form of the organs and their development—this will be treated in a series of articles dealing with the great subdivisions of plants (see Angiosperms, Gymnosperms, Pteridophyta, Bryophyta, Algae, Lichens, Fungi and Bacteriology) and the more important organs (see Stem, Leaf, Root, Flower, Fruit); (b) Anatomy, the study of internal structure, including minute anatomy or histology (see Plants: Anatomy).

2\. Cytology (q.v.), the intimate structure and behaviour of the cell and its contents—protoplasm, nucleus, &c.

3\. Physiology, the study of the life-functions of the entire plant and its organs (see Plants: Physiology).

4\. Systematic, the arrangement and classification of plants (see Plants: Classification).

5\. Distribution or Geographical Botany, the consideration of the distribution of plants on the earth’s surface (see Plants: Distribution).

6\. Palaeontology, the study of the fossils found in the various strata of which the earth is composed (see Palaeobotany).

7\. Ecology or Oecology, the study of plants in relation to each other and to their environment (see Plants: Ecology).

Besides these departments which deal with Botany as a science, there are various applications of botany, such as forestry (see Forests and Forestry), agriculture (q.v.), horticulture (q.v.), and materia medica (for use in medicine; see the separate articles on each plant).

(A. B. R.)

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1 Morison, Pradudia Botanica (1672); Plantarum Historia Universalis (1680).

2 Rivinus (Augustus Quirinus) paterno nomine Bachmann, Introductio genetatis in Rem Herbariam (Lipsiae, 1690).

3 Tournefort, Élémens de botanique (1694); Institutiones Rei Herbariae (1700).

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