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History of Plant Sciences

Janet Browne, Wellcome Trust Centre for the History of Medicine, London, UK
The study of plants has played a significant, if sometimes undervalued, role in history. Plants have been important in science, medicine and economic affairs since Antiquity. The herbal tradition dominated investigations until the Renaissance after which it ran alongside developing areas of expertise in classification and physiology. In the eighteenth century, plant sciences became significant in geographical exploration and in general culture. Nineteenth-century botanists used plants for investigating cell theory and early genetics as well as in evolutionary biology.

Introductory article
Article Contents
. Introduction . Early Ideas . Expanding Horizons . Plant Science in Nineteenth-century Thought

Introduction
The scientific study of plants today is usually distinguished from endeavours in which plants are used for a particular purpose, as in economic botany, horticulture, agriculture, materia medica and the like. But this was not the case in the past. From time immemorial people have studied plants precisely for the uses they may have for human beings, and this tradition has been inseparable from the development of western botanical science. The interrelationships still hold in the case of herbal medicine, for example, especially in the search for alternative remedies to manufactured drugs and in the practices of ethnobotany. The modern sciences of plant genetics depend extensively on horticultural and agricultural input. Commercial development of plantation crops and global trade in plant-based products, such as cotton, play key roles in national economies. Because of these longlasting links with human activities and economic concerns, botany has often been called the ‘big science’ of previous centuries.

preparing charcoal and obtaining pitch from resinous trees. By far the most famous Ancient text was by Dioscorides (fl. ad 50–70). Dioscorides’ herbal, a plant encyclopedia written in Greek and intended for the use of physicians, was translated into Latin and Arabic, and copied and recopied incessantly, often with additions and manuscript illustrations. Above all, herbalists and apothecaries had to be sure of the plant’s identity. Consequently written descriptions became lengthy and illustrations important. The advent of printing around 1453 accelerated this trend and Dioscorides’ herbal was one of the first texts to be produced by this method in 1478. The book became the foundation of many of the greatest illustrated herbals, particularly that of Otto Brunfels (c. 1489–1534), who drew illustrations directly from the plants, and Leonhart Fuchs (1501–1566), who produced a magnificent herbal, De Historia Stirpium, in 1542, figuring about 550 plants. These were expensive books, often hand-coloured and annotated. In this regard, learned botany was restricted mainly to physicians, theologians, university dons and the court.

Early Ideas
Much of the earliest botany was concerned with materia medica. An exception was the work of the Ancient Greek philosopher Theophrastus (c. 371–287 bc) who studied plants in detail, although his work was not known in the west until the Renaissance. Theophrastus was Aristotle’s favourite pupil and successor at the Lyceum. In essence he did for plants what Aristotle had done for animals. He classified plants in the Aristotelian tradition, expounding the basic formal principles of taxonomy using a logical division of genera and species to yield a definition of one natural kind or form. The definition applied to that one form alone. On the larger scale, he divided plants into the categories tree, shrub, under-shrub and herb. In addition, he gave his observations on reproduction and nutrition, correctly identifying the need for pollination between male and female date palms. He included much information on the agricultural techniques of his day such as methods of

Expanding Horizons
After the fall of Rome, Greek and Roman knowledge was preserved and extensively improved in Arabic translation. The revival of these Ancient texts recast western botany in the later sixteenth century. Practitioners increasingly noticed that the plants of western Europe were not those described in either Arabic or Ancient texts and began to add to traditional lists. This coincided with increasing foreign travel, the colonial expansion of Europe, most notably into the East Indies and the New World, and changing medical education, itself partly a consequence of the new texts available. An increasing number of exotic plants and herbs were introduced to Europe from overseas, encouraging the development of the small physic gardens already attached to the main medical universities, enrich1

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writing in English and Latin respectively. John Ray (1627–1705) notably pushed theories of classification forward. He seems to have seen spores in ferns. sepals and reproductive organs. nerves or respiratory system. ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.and seventeenth-century science in general. at the Royal Society.net . By the end of the century Rembert Dodoens (1516–1585) and William Turner (1508–1568) were producing works that were not so much supplements to the Ancients as true replacements. The chemical composition of various plants was established. He used the word ‘staticks’ to convey his emphasis on precise measurement and was one of several experimenters of the period to see the value of using controls in his experiments.History of Plant Sciences ing the gardens of the royal courts. hygiene and dietetics. The first controlled experiments into plant nutrition were performed by J. Using relatively simple microscopes. for it appealed to an increasingly wider. He also produced an important catalogue of English plants. was in plant anatomy. He conclusively showed that sap did not circulate like blood circulates in animals. The botanical treatises of Gaspard Bauhin (1560–1624) and the French botanist de L’Ecluse (Latinized to Clusius. finding that the tree had gained some 170 lb from water alone. who produced 11 editions of the English Physician Enlarged. B. should be used to divide plants hierarchically into ‘natural’ groups. he weighed the tree and the annual leaf fall. Each author attempted his own solution to the problem of how much weight to give to various botanical criteria. and each searched for a ‘natural’ arrangement of kinds – a classification scheme that would reflect morphological relationships (or ‘affinities’) as seen in the number and relative position of petals. and Marcello Malpighi (1628–1694). During the major intellectual changes that took place in the seventeenth century. the monocotyledons and dicotyledons. van Helmont (1579–1644) in the 1640s. especially the reproductive parts. he investigated the movement of sap and water in the plant. Bauhin’s Pinax of 1623 described some 6000 plants. and serve as opening points to the great age of classical taxonomy. and showed that similar structures occur in many plants. Cesalpino returned to Aristotle’s and Theophrastus’ logical hierarchies and gave a series of ‘essential’ characters whose successive subdivisions would not violate intuitive perceptions of affinity. for instance finding that flowering plants fall into two natural groups based on the number of seedleaves (cotyledons). The work of Andrea Cesalpino (1519–1603) at the University of Pisa was a landmark in classification schemes. perceived as the condensed growing point of a shoot. giving descriptions of useful plants. and followed the cycle of development from seed to seed.els. although the question remained hotly contested for decades afterwards. and posing questions about geographical distribution. mosses and two kinds of mould. He performed experiments on the gas relations of plants. Grew proposed that the stamens were the male reproductive organ and that plants possessed two sexes. He decided that the structures concerned with the plant’s primary functions. Meanwhile. Some of their most significant observations were on the origin of buds. 2 Similarly. A few years later Stephen Hales (1677–1761) began careful experimental studies of plant physiology. so that for Grew the large xylem vessels seemed as if they might function as air-vessels like the tracheae of animals. in Bologna. Nomenclature was much advanced by Joseph Pitton de Tournefort (1656–1708) when he defined and ´ le´ments de Botanique in 1694. The Royal Society in London was one of the prime locations for such research. or the Herbal (1653). the field of botany broadened to come into contact with other disciplines besides medicine and classification. including stem. Nehemiah Grew (1641–1712). The functions of such structures were mostly interpreted by analogy with animals. which he called ‘cells’. more fundamental divisions of the plant kingdom. rather in the way that animal classification schemes could use differences in the heart. more popular audience. He weighed and planted a willow tree in a container and allowed it to grow for five years. the shape of leaves. published in Vegetable Staticks (1727). Italy and The Netherlands revealed the unsuspected world of the small. root and leaf. and recipes for simple medications. He was especially gifted in defining the higher. albeit slowly and not as dramatically as contemporary advances in understanding animal physiology. Herbals continued to be published in large numbers in relatively stereotyped format. Attempts to order and classify natural items were primary activities in sixteenth. and means of reproduction. whether by bulbs or seeds. Proof of this suggestion came in the work of Rudolf Jacob Camerarius in 1694. in order to throw light on the chemical composition of water. however. at the end. 1526–1609) were exemplary in this regard. where the use of the microscope by skilled natural philosophers in Britain. Cesalpino’s scheme underlay most of the more important natural plant classifications published afterwards. Nicholas Culpeper (1616–1654). they identified the principal tissues of the plant body. as part of the path-breaking studies on ‘airs’ performed by Robert Boyle and Joseph Priestley. led this movement. again like animals. the herbal tradition was scarcely affected by these philosophical inquiries. independently developed an integrated view of microscopic plant structure. The active experimental inquiries that characterized other natural philosophical areas penetrated botany too. Taking his cue from animal physiology. their habitats and properties. His chief achievement was to establish the constant uptake of water by plants and its loss by transpiration (‘perspiration’ he called it) but he also showed that these movements took place by root and leaf suction. Robert Hooke (1635–1703) saw ‘pores’ in cork. named nearly 700 genera in E The most immediate extension of botanical investigation. Later herbals included advice on domestic economy. representing the rapid increase in knowledge over the previous hundred years.

Hothouses and stove-plants became an increasing possibility for the wealthy. for example. such as the growing interest in landscape gardening and the cult of the picturesque. The most famous eighteenth-century naturalist. whom he called ‘apostles’. cheap editions. and sometimes criticizing. the Carolinas. commercial entrepreneurs and government officials jointly opened up these routes to economic expansion. by subsequent convention. the botanist James Edward Smith (1759–1828) purchased his collections. Previously. and the great European gardens such as the Jardin des Plantes in Paris. the Cape. central Spain. as well as beautifully coloured illustrated works for richer clientele. There. South Africa. Linnaeus also helped botanical knowledge to move out of the elite world of universities. which profoundly changed European ideas about botanical science. After the death of Linnaeus in 1778. under Sir Joseph Banks. his colleagues by naming plants after them. came through geographical expeditions. paralleling similar desires to provide classification schemes in chemistry and medicine. Travellers and collectors could quickly allocate unidentified new specimens into a class or family. the first name denoting the genus. Many of these exotics came from government voyages of exploration to Australia.History of Plant Sciences Through these volumes. Linnaeus proposed an ‘artificial system’ for plants whereby the numbers of stamens and pistils (male and female reproductive organs) were counted and served to allocate plants into groups without any further ado. definitions of a species could run to a sentence or more because so many key criteria needed to be included. initiating a rage for choice specimens and providing the foundations of national herbaria and museum collections. Nevertheless. In actual fact Linnaeus did regard the reproductive system as having primary physiological importance and his scheme was not intended to be solely arbitrary. for it grouped plants together on the basis of numbers alone. who often possessed illustrated floras and synopses of classification schemes in their libraries. and the South Seas. was usually understood as a method of classifying plants purely for human convenience. the ´ (Latinized as Carolus Swedish doctor. In his Genera Plantarum and Species Plantarum he defined all the known plants of the world. Plants played a significant role in the life of the landed elite. and gave rise to some curious juxtapositions. These men. transformed botanical science in all quarters. been taken as the type. countless people learned a utilitarian and domestic combination of medical and botanical science. Living plants could be seen at Kew Gardens (royal property until 1841). museums and physic gardens into a far broader constituency. He introduced the convention of binomial names. the second the specific. Women were particularly prominent as authors of elementary textbooks. one of Linnaeus’s favoured pupils. became a hub of proto-imperial science. European prosperity overseas equally depended on the development of the plantation system in which staple crops like tea or sugar-cane were relocated for colonial purposes. was collected and first classified by Linnaeus’s method. are all known to have first encountered botany through popularizations of Linnaeus’s classification scheme. Many eighteenthcentury people in Europe and North America. and explanations were published. In a succession of philosophical writings Linnaeus set out principles for the science. or Syon Park near Chiswick. Sir Joseph Banks accompanied James Cook on his first voyage in search of the Southern continent (Australia). however. The landed gentry cultivated exotic plants and patronized plant collectors and landscape gardeners. and so on. Enterprising private societies like the Royal Horticultural Society (founded in 1804) sponsored collecting trips abroad and ran public gardens and competitions for their members.els. Many popular books. In particular. Botany’s greatest impact. sent numerous collections back to Europe throughout the eighteenth century. Naval officers. These names and definitions have. Eighteenth-century botany therefore had wide cultural appeal. They achieved remarkable results. since botany was widely perceived as a suitable science for genteel women to pursue. In Britain. This system. Other aspects of botanical science should not be ignored either. ranging from sexual innuendo about plants as humans. Asia. He personally coined many of the names still in use and enjoyed complimenting. 3 ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www. for example plants with five stamens would be placed in a new category called Pentandria (Greek for five male organs). Carl von Linne Linnaeus. He provided a quick and easy alternative to existing plant classification schemes in his Systema naturae (1735). and others using his system. it was simple to count stamens: anyone could do it.net . women and working men. sometimes called the numerical system or the sexual system. the Linnaean system encouraged a number of women to participate in science as authors. translations. and Kew Gardens. Indeed botany was one of the few sciences open to women. including gentlefolk. In 1772. taking with him Daniel Solander (1733–1782). He trained and sent plant collectors all over the increasingly accessible globe: to Japan. or in an anti-Catholic classification of monks. In this sense Linnaeus’s scheme liberated many of those people who worked with plants. Chatsworth. The magnificent collection of plants brought back by Banks and Solander. and Linnaeus consciously acted as the hub of a vast botanical exploring network fanning out from his base at the medical school of Uppsala University. His ‘sexual system’ provided the basis for much cultural satire and parody. These introduced a large number of new species to the west. His impact stretched into all realms of scientific thought. the East India Company took the lead in establishing botanic gardens in Saharanpore and Calcutta. 1707–1778).

modification and fusion. Auguste de Candolle formulated useful concepts of symmetry. who concluded that the pollen tube conveys ‘spermatic fluid’ to the ovule. Flora Britannica (1800– 1804). He confirmed Cuvier’s view that the living beings of the Earth had changed in character in each successive geological era. abortion. Smith established the Linnean Society of London in 1788 (the society’s name is ´ . The natural system came to be understood as a means of classification in which a number of morphological characters and charactercomplexes were used to determine relationships. where he delivered an address entitled ‘Introductory Discourse on the Rise and Progress of Natural History’. reproduction and anatomy. uninhabitable state to one in which the present state of affairs emerged. But fossil plants were not really integrated into geological thought until the issue of the central heat of the Earth arose in the closing years of the eighteenth century. Plant Science in Nineteenth-century Thought In the opening decades of the nineteenth century Robert Brown greatly advanced the use of the microscope in plant embryological investigations. Towards the end of the eighteenth century growing knowledge of plant fossils helped naturalists consider the course of the Earth’s history. Fossil plants had been noted and displayed in collectors’ cabinets since the time of Edward Lhuyd (1660–1709) and were pictured in some detail by J. although here the major explanatory theories were principally derived from structural geology and animal evidence. Smith was elected president at the first meeting. 4 The study of palaeobotany came into its own with the work of Adolphe Theodore Brongniart (1801–1876). A unique contribution to botany was made by Johann Wolfgang Goethe (1749–1832). little obvious attention was paid to plant palaeontology until the rise of Quarternary studies in the twentieth century. However. Scheuchzer understood plant fossils to be the remains of species killed by Noah’s Flood. The Swiss botanist Auguste de Candolle (also known as Augustin Pyramus de Candolle. buried in obscure articles or in systematic works. This was not an entirely original concept. He later published English Botany (1790) and his most important work. foliage leaves and the floral parts. Beyond this. Die Metamorphose der Pflanzen (1790) (translated into English in 1863 as Essay on the Metamorphosis of Plants). Later. Scheuchzer (1672–1733) in his Herbarium Diluvianum of 1709. for the most part.net . 1778–1841) expanded his interpretation by studying coal deposits in more detail. Candolle proposed that the preponderance of cycad and fern remains in coal indicated that the early atmosphere of the Earth must have been very much warmer and possibly richer in carbon dioxide. Smaller societies and specimen exchange clubs also proliferated. Goethe was the first to express what became a much-debated issue in plant morphology. Jean Baptista Amici (1786–1863) observed the pollen tube growing out of pollen grains lodged on the stigmatic surface. and is spelled without taken from von Linne the first ‘a’). Much of what Adanson proposed has been retained in today’s broad-based taxonomical schemes. Even Linnaeus himself conceded at the end of his life that some hybridization must occur and that perhaps a few of the major plant groups had originated in that way: one ‘mother’ group hybridizing with several ‘fathers’ to produce related botanical families. in fact. In 1827 he described the development of the unfertilized ovule. and the action of pollen in fertilization helped in understanding animal reproduction. naturalists vigorously investigated physiology. The nature of coal deposits was. In Florence. In specialist circles. allowing him to draw a fundamental distinction between gymnosperms (conifers and cycads) and angiosperms (flowering plants). This helped Brown decide that the ovule in cycads. Other alternatives to rigidly logical classification schemes emerged as a number of botanists searched for ways to acknowledge a wider range of resemblances. readily drawing parallels between animals and plants. J. that in essence all floral parts were modified leaves. Antoine de Jussieu (1686–1758) observed that the plants in the coal deposits around Lyon were typical of the hot and humid climates of the equator. Questions about the definition of life and organization were asked. was based on the idea of homology between plant organs such as cotyledons. Brown (1773–1858) made many brilliant observations in microscopic plant anatomy that were. a leading issue in geological circles although not much connected with commercial enterprise. Michel Adanson (1727–1806) in his important Famille des Plantes (1763) attempted to work out purely empirical relationships by including a large number of characters. published as Histoire des Ve´ge´taux Fossiles (1828). fixed since their creation by God. In 1831 Brown then delivered a detailed account ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www. most of which used the Linnaean system of classification. stimulated studies of hybrization. not Linnaeus. These observations were repeated and confirmed by Adolphe Brongniart.History of Plant Sciences originally offered to Joseph Banks. Sensitive plants like the mimosa were thought to possess animal-like qualities such as irritability. He identified more fossil plants than ever before and used them to reconstruct the physical history of the Earth. Linnaeus’s definition of plant species as stable entities. conifers and other related families is truly naked and not enclosed at any stage in an ovary. Studies of ‘breathing’ in plants played a significant role in the discovery of oxygen and then photosynthesis. which soon became the premier botanical society in England although it ostensibly dealt with all branches of the living world. This view contributed to contemporary theories about the directional history of the Earth from a hot.els.

Kew. Although he could not explain the cause of osmosis. Frederick Bower (1855–1948) linked the migration of plants onto land to the appearance and subsequent elaboration of an asexual generation. Subsequently an important step in plant physiology came from Henri Dutrochet (1776–1847). Celakowsky saw the evolutionary significance of Hofmeister’s work. His results appeared in 1837. including the transport of seeds by birds and currents. the emphasis of university botany shifted so much towards the physiological. and Asa Gray (1818– 1888). Botanists probably found the idea of plant evolution far less repugnant than that of human evolution. The idea of the alternation of generations in animals helped Wilhelm Hofmeister (1824–1877) demonstrate the two phases in the life cycle of mosses and liverworts in 1851. By Mohl’s time. Later he worked extensively on the cross-fertilization of plants. and did not worry too much about the moral and theological implications of plant descent. Darwin also performed plant experiments to investigate the origin of variations. Brown’s work on cell anatomy was quickly followed by Mathias Schleiden’s path-breaking Grundzuge der wissenschaftlichen Botanik (1842) translated into English in 1849 as Principles of Scientific Botany. and then Glasgow in 1885 revolutionized the way plant physiology was regarded. Among other achievements. Both disciplines were founded on a descriptive anatomical tradition. where free-swimming spermatozoids were identified. director of the Royal Botanic Gardens. for example in enzyme 5 ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.History of Plant Sciences of the process of fertilization in orchids and Asclepiadeae. Even so. Boussingault (1822–1895). Darwin studied orchids as a prime example of adaptation. Nevertheless plants have often seemed secondary in evolutionary theory even though botany dominated Charles Darwin’s (1809–1882) later researches. vague notions of utricles and vessels had given way to a clear concept of the plant cell based on a firm body of knowledge about its structure. Many of the advances made during these years were summarized by Hugo von Mohl (1805–1872) in 1851.net . and tropisms (plant hormones). Sachs made highly significant studies on plant hormones. polymorphism and hybrids. in this regard as adaptations to ensure insect fertilization. Professor of Botany at Harvard University. Hofmeister was followed by Ladislav Celakowsky who in 1868 coined the terms gametophyte and sporophyte. Thereafter Brown was renowned for noting the form of the cell nucleus. Experimental botany at first failed to find a secure home in the English universities. His two chapters on geographical distribution presented innovative ideas about historical plant geography. and worked on pollination. Throughout the nineteenth century Engler’s arrangement was enshrined in most classification schemes until C. After the Origin of Species. such as the effect of gravity or light on roots and shoots.els. Elsewhere fundamental observations were made on the nature of fertilization in algae. he did not doubt that it was an entirely physical process and was of very great importance in vital phenomena. The nitrogen cycle was elucidated by J. arguing from the phylogenetic perspective that sexual generation must be more primitive than asexual. who studied medicine in Paris but devoted much of his life to the scientific investigation of osmosis. In the Origin of Species (1859) Darwin used botanical investigations to support his key arguments for natural selection. he focused on the active physiological life of the organism. In 1868 the first edition of Julius von Sachs’ Lehrbuch der Botanik appeared which led the way in plant physiological investigations in Europe. on insectivorous plants and the movements of plants. both imported the new laboratory work wholesale from the continent. Heated debate arose over the origin of flowering plants (angiosperms) in the Cretaceous period. where he discussed the possibility of northern species pushing through the tropics at some former cool period. J. This also involved controversy over what should properly be considered primitive. The morphology of plants was generally underrated until evolutionary theory gave it new legitimacy. In all this work. Gray and Hooker were among the first to use natural selection for understanding plant relationships. His close friends Joseph Hooker (1817–1911). mode of differentiation and even some of its chemical constituents. there were a number of satirical references and cartoons poking fun at the transformation of turnips into humans. and the means of plant dispersal. and both found it hard to become separate from the medical curriculum. Sachs’ impact was as great as Linnaeus’s some hundred years before although in a very different field. proposed that Ranunculus and its relatives came closest among living plants to primitive angiosperms. In this work Schleiden (1804–1881) laid the groundwork for the cell theory of Theodor Schwann. however. Bower’s appointments in London. in 1915. E. the evolution of sexes. Every eminent plant physiologist was trained in Sachs’ laboratory in Wurzburg. reproduction. Generally. D. with the exception of John Burdon Sanderson at University College London and Sidney Vines in Cambridge. Bessey. Elsewhere. showing how plants vary in the wild and display adaptations. In many ways the state of English botany before then resembled that of English physiology. there were few fights over the evolution of plants in the same way as there were fights over the possible connections between animals and humankind. growth mechanisms and tropisms. B. Modern perceptions of the overwhelming importance of evolutionary theory in the nineteenth century have obscured these dramatic surges in physiological and microscopical investigations. The essential steps in carbon fixation were not recognized until Julius von Sachs (1832–1897) showed that the starch present in green cells was the product of the carbon dioxide absorbed. were both converted to evolutionary theory and defended Darwin’s ideas in reviews and in their botanical writings.

Delaporte F (1982) Nature’s Second Kingdom.net . Explorations of Vegetality in the Eighteenth Century. Frey KJ (ed. The Key to Natural Selection. and cytology. Goldhammer A (transl. Green JR (1909) A History of Botany. Ithaca. Oxford: Clarendon Press. 6 ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www. Further Reading Allan MA (1977) Darwin and his Flowers. Ames: Iowa State University Press. Andrews HN (1980) The Fossil Hunters: In Search of Ancient Plants. London: Faber and Faber. Cambridge. fertilization and cell division.History of Plant Sciences studies and fermentation. that Hooker felt it necessary to reassert the value of taxonomy as an academic discipline in the Index Kewensis and his and Bentham’s Genera Plantarum. Botany went on to become the prime area for genetic research. Being a Continuation of Sachs’ History of Botany 1530 – 1860.) (1994) Historical Perspectives in Plant Science. NY: Cornell University Press.els. as in pure line experiments.). MA: MIT Press. London: Academic Press. Morton AG (1981) History of Botanical Science. 1860 – 1900. An Account of the Development of Botany from Ancient Times to the Present Day.