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 scientic 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 (. ad 5070). 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 plants 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 rst 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. 14891534), who drew illustrations directly from the plants, and Leonhart Fuchs (15011566), who produced a magnicent herbal, De Historia Stirpium, in 1542, guring 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. 371287 bc) who studied plants in detail, although his work was not known in the west until the Renaissance. Theophrastus was Aristotles favourite pupil and successor at the Lyceum. In essence he did for plants what Aristotle had done for animals. He classied plants in the Aristotelian tradition, expounding the basic formal principles of taxonomy using a logical division of genera and species to yield a denition of one natural kind or form. The denition 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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History of Plant Sciences

ing the gardens of the royal courts, and posing questions about geographical distribution. By the end of the century Rembert Dodoens (15161585) and William Turner (15081568) were producing works that were not so much supplements to the Ancients as true replacements. Attempts to order and classify natural items were primary activities in sixteenth- and seventeenth-century science in general. The botanical treatises of Gaspard Bauhin (15601624) and the French botanist de LEcluse (Latinized to Clusius, 15261609) were exemplary in this regard, and serve as opening points to the great age of classical taxonomy. 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 classication scheme that would reect morphological relationships (or anities) as seen in the number and relative position of petals, sepals and reproductive organs, the shape of leaves, and means of reproduction, whether by bulbs or seeds. Bauhins Pinax of 1623 described some 6000 plants, representing the rapid increase in knowledge over the previous hundred years. The work of Andrea Cesalpino (15191603) at the University of Pisa was a landmark in classication schemes. Cesalpino returned to Aristotles and Theophrastus logical hierarchies and gave a series of essential characters whose successive subdivisions would not violate intuitive perceptions of anity, rather in the way that animal classication schemes could use dierences in the heart, nerves or respiratory system. He decided that the structures concerned with the plants primary functions, especially the reproductive parts, should be used to divide plants hierarchically into natural groups. Cesalpinos scheme underlay most of the more important natural plant classications published afterwards. During the major intellectual changes that took place in the seventeenth century, the eld of botany broadened to come into contact with other disciplines besides medicine and classication. John Ray (16271705) notably pushed theories of classication forward. He was especially gifted in dening the higher, more fundamental divisions of the plant kingdom, for instance nding that owering plants fall into two natural groups based on the number of seedleaves (cotyledons), the monocotyledons and dicotyledons. He also produced an important catalogue of English plants. Nomenclature was much advanced by Joseph Pitton de Tournefort (16561708) when he dened and lements de Botanique in 1694. named nearly 700 genera in E The most immediate extension of botanical investigation, however, was in plant anatomy, where the use of the microscope by skilled natural philosophers in Britain, Italy and The Netherlands revealed the unsuspected world of the small. The Royal Society in London was one of the prime locations for such research. Robert Hooke (16351703) saw pores in cork, which he called cells, and showed that similar structures occur in many plants. He seems to have seen spores in ferns, mosses and two kinds of mould.
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Similarly, Nehemiah Grew (16411712), at the Royal Society, and Marcello Malpighi (16281694), in Bologna, writing in English and Latin respectively, independently developed an integrated view of microscopic plant structure. Using relatively simple microscopes, they identied the principal tissues of the plant body, including stem, root and leaf, and followed the cycle of development from seed to seed. Some of their most signicant observations were on the origin of buds, perceived as the condensed growing point of a shoot. The functions of such structures were mostly interpreted by analogy with animals, so that for Grew the large xylem vessels seemed as if they might function as air-vessels like the tracheae of animals. Grew proposed that the stamens were the male reproductive organ and that plants possessed two sexes, again like animals. Proof of this suggestion came in the work of Rudolf Jacob Camerarius in 1694, although the question remained hotly contested for decades afterwards. The active experimental inquiries that characterized other natural philosophical areas penetrated botany too, albeit slowly and not as dramatically as contemporary advances in understanding animal physiology. The rst controlled experiments into plant nutrition were performed by J. B. van Helmont (15791644) in the 1640s, in order to throw light on the chemical composition of water. He weighed and planted a willow tree in a container and allowed it to grow for ve years; at the end, he weighed the tree and the annual leaf fall, nding that the tree had gained some 170 lb from water alone. The chemical composition of various plants was established. A few years later Stephen Hales (16771761) began careful experimental studies of plant physiology, published in Vegetable Staticks (1727). 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. Taking his cue from animal physiology, he investigated the movement of sap and water in the plant. 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. He conclusively showed that sap did not circulate like blood circulates in animals. He performed experiments on the gas relations of plants, as part of the path-breaking studies on airs performed by Robert Boyle and Joseph Priestley. Meanwhile, the herbal tradition was scarcely aected by these philosophical inquiries, for it appealed to an increasingly wider, more popular audience. Nicholas Culpeper (16161654), who produced 11 editions of the English Physician Enlarged, or the Herbal (1653), led this movement. Herbals continued to be published in large numbers in relatively stereotyped format, giving descriptions of useful plants, their habitats and properties, and recipes for simple medications. Later herbals included advice on domestic economy, hygiene and dietetics.

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History of Plant Sciences

Through these volumes, countless people learned a utilitarian and domestic combination of medical and botanical science. The most famous eighteenth-century naturalist, the (Latinized as Carolus Swedish doctor, Carl von Linne Linnaeus, 17071778), transformed botanical science in all quarters. He provided a quick and easy alternative to existing plant classication schemes in his Systema naturae (1735). There, Linnaeus proposed an articial 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, for example plants with ve stamens would be placed in a new category called Pentandria (Greek for ve male organs). This system, sometimes called the numerical system or the sexual system, was usually understood as a method of classifying plants purely for human convenience, for it grouped plants together on the basis of numbers alone; and gave rise to some curious juxtapositions. In actual fact Linnaeus did regard the reproductive system as having primary physiological importance and his scheme was not intended to be solely arbitrary. Nevertheless, it was simple to count stamens: anyone could do it. In this sense Linnaeuss scheme liberated many of those people who worked with plants. Travellers and collectors could quickly allocate unidentied new specimens into a class or family; 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. He trained and sent plant collectors all over the increasingly accessible globe: to Japan, South Africa, the Carolinas, Asia, central Spain, and so on. These men, whom he called apostles, and others using his system, sent numerous collections back to Europe throughout the eighteenth century. They achieved remarkable results. In 1772, for example, Sir Joseph Banks accompanied James Cook on his rst voyage in search of the Southern continent (Australia), taking with him Daniel Solander (17331782), one of Linnaeuss favoured pupils. The magnicent collection of plants brought back by Banks and Solander, which profoundly changed European ideas about botanical science, was collected and rst classied by Linnaeuss method. In a succession of philosophical writings Linnaeus set out principles for the science. He introduced the convention of binomial names, the rst name denoting the genus, the second the specic. Previously, denitions of a species could run to a sentence or more because so many key criteria needed to be included. In his Genera Plantarum and Species Plantarum he dened all the known plants of the world. These names and denitions have, by subsequent convention, been taken as the type. He personally coined many of the names still in use and enjoyed complimenting, and sometimes criticizing, his colleagues by naming plants after them.

Linnaeus also helped botanical knowledge to move out of the elite world of universities, museums and physic gardens into a far broader constituency. Many eighteenthcentury people in Europe and North America, including gentlefolk, women and working men, are all known to have rst encountered botany through popularizations of Linnaeuss classication scheme. Many popular books, translations, cheap editions, and explanations were published, as well as beautifully coloured illustrated works for richer clientele. His impact stretched into all realms of scientic thought, paralleling similar desires to provide classication schemes in chemistry and medicine. In particular, the Linnaean system encouraged a number of women to participate in science as authors. His sexual system provided the basis for much cultural satire and parody, ranging from sexual innuendo about plants as humans, or in an anti-Catholic classication of monks. Eighteenth-century botany therefore had wide cultural appeal. Other aspects of botanical science should not be ignored either, such as the growing interest in landscape gardening and the cult of the picturesque. Plants played a signicant role in the life of the landed elite, who often possessed illustrated oras and synopses of classication schemes in their libraries. The landed gentry cultivated exotic plants and patronized plant collectors and landscape gardeners. Hothouses and stove-plants became an increasing possibility for the wealthy. Women were particularly prominent as authors of elementary textbooks, since botany was widely perceived as a suitable science for genteel women to pursue. Indeed botany was one of the few sciences open to women. Botanys greatest impact, however, came through geographical expeditions. These introduced a large number of new species to the west, initiating a rage for choice specimens and providing the foundations of national herbaria and museum collections. Many of these exotics came from government voyages of exploration to Australia, the Cape, and the South Seas. Living plants could be seen at Kew Gardens (royal property until 1841), Chatsworth, or Syon Park near Chiswick, and the great European gardens such as the Jardin des Plantes in Paris. Enterprising private societies like the Royal Horticultural Society (founded in 1804) sponsored collecting trips abroad and ran public gardens and competitions for their members. 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. Naval ocers, commercial entrepreneurs and government ocials jointly opened up these routes to economic expansion. In Britain, the East India Company took the lead in establishing botanic gardens in Saharanpore and Calcutta; and Kew Gardens, under Sir Joseph Banks, became a hub of proto-imperial science. After the death of Linnaeus in 1778, the botanist James Edward Smith (17591828) purchased his collections,
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History of Plant Sciences

originally oered to Joseph Banks. Smith established the Linnean Society of London in 1788 (the societys name is , not Linnaeus, and is spelled without taken from von Linne the rst a), which soon became the premier botanical society in England although it ostensibly dealt with all branches of the living world. Smith was elected president at the rst meeting, where he delivered an address entitled Introductory Discourse on the Rise and Progress of Natural History. He later published English Botany (1790) and his most important work, Flora Britannica (1800 1804). Smaller societies and specimen exchange clubs also proliferated, most of which used the Linnaean system of classication. In specialist circles, naturalists vigorously investigated physiology, reproduction and anatomy, readily drawing parallels between animals and plants. Sensitive plants like the mimosa were thought to possess animal-like qualities such as irritability. Studies of breathing in plants played a signicant role in the discovery of oxygen and then photosynthesis; and the action of pollen in fertilization helped in understanding animal reproduction. Linnaeuss denition of plant species as stable entities, xed since their creation by God, stimulated studies of hybrization. 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. Questions about the denition of life and organization were asked. Other alternatives to rigidly logical classication schemes emerged as a number of botanists searched for ways to acknowledge a wider range of resemblances. Michel Adanson (17271806) in his important Famille des Plantes (1763) attempted to work out purely empirical relationships by including a large number of characters. Much of what Adanson proposed has been retained in todays broad-based taxonomical schemes. The natural system came to be understood as a means of classication in which a number of morphological characters and charactercomplexes were used to determine relationships. Towards the end of the eighteenth century growing knowledge of plant fossils helped naturalists consider the course of the Earths history, although here the major explanatory theories were principally derived from structural geology and animal evidence. Fossil plants had been noted and displayed in collectors cabinets since the time of Edward Lhuyd (16601709) and were pictured in some detail by J. J. Scheuchzer (16721733) in his Herbarium Diluvianum of 1709. Scheuchzer understood plant fossils to be the remains of species killed by Noahs Flood. 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. Antoine de Jussieu (16861758) observed that the plants in the coal deposits around Lyon were typical of the hot and humid climates of the equator.
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The study of palaeobotany came into its own with the work of Adolphe Theodore Brongniart (18011876). He identied more fossil plants than ever before and used them to reconstruct the physical history of the Earth, published as Histoire des Vegetaux Fossiles (1828). He conrmed Cuviers view that the living beings of the Earth had changed in character in each successive geological era. The Swiss botanist Auguste de Candolle (also known as Augustin Pyramus de Candolle, 17781841) 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. This view contributed to contemporary theories about the directional history of the Earth from a hot, uninhabitable state to one in which the present state of aairs emerged. The nature of coal deposits was, in fact, a leading issue in geological circles although not much connected with commercial enterprise. Beyond this, little obvious attention was paid to plant palaeontology until the rise of Quarternary studies in the twentieth century. A unique contribution to botany was made by Johann Wolfgang Goethe (17491832). Die Metamorphose der Panzen (1790) (translated into English in 1863 as Essay on the Metamorphosis of Plants), was based on the idea of homology between plant organs such as cotyledons, foliage leaves and the oral parts. This was not an entirely original concept. However, Goethe was the rst to express what became a much-debated issue in plant morphology, that in essence all oral parts were modied leaves. Later, Auguste de Candolle formulated useful concepts of symmetry, abortion, modication and fusion.

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. Brown (17731858) made many brilliant observations in microscopic plant anatomy that were, for the most part, buried in obscure articles or in systematic works. In 1827 he described the development of the unfertilized ovule. This helped Brown decide that the ovule in cycads, conifers and other related families is truly naked and not enclosed at any stage in an ovary, allowing him to draw a fundamental distinction between gymnosperms (conifers and cycads) and angiosperms (owering plants). In Florence, Jean Baptista Amici (17861863) observed the pollen tube growing out of pollen grains lodged on the stigmatic surface. These observations were repeated and conrmed by Adolphe Brongniart, who concluded that the pollen tube conveys spermatic uid to the ovule. In 1831 Brown then delivered a detailed account

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of the process of fertilization in orchids and Asclepiadeae. Thereafter Brown was renowned for noting the form of the cell nucleus. Browns work on cell anatomy was quickly followed by Mathias Schleidens path-breaking Grundzuge der wissenschaftlichen Botanik (1842) translated into English in 1849 as Principles of Scientic Botany. In this work Schleiden (18041881) laid the groundwork for the cell theory of Theodor Schwann. Many of the advances made during these years were summarized by Hugo von Mohl (18051872) in 1851. By Mohls time, vague notions of utricles and vessels had given way to a clear concept of the plant cell based on a rm body of knowledge about its structure, reproduction, mode of dierentiation and even some of its chemical constituents. Elsewhere fundamental observations were made on the nature of fertilization in algae, where free-swimming spermatozoids were identied. Subsequently an important step in plant physiology came from Henri Dutrochet (17761847), who studied medicine in Paris but devoted much of his life to the scientic investigation of osmosis. His results appeared in 1837. Although he could not explain the cause of osmosis, 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. B. J. D. Boussingault (18221895). The essential steps in carbon xation were not recognized until Julius von Sachs (18321897) showed that the starch present in green cells was the product of the carbon dioxide absorbed. Modern perceptions of the overwhelming importance of evolutionary theory in the nineteenth century have obscured these dramatic surges in physiological and microscopical investigations. Nevertheless plants have often seemed secondary in evolutionary theory even though botany dominated Charles Darwins (18091882) later researches. In the Origin of Species (1859) Darwin used botanical investigations to support his key arguments for natural selection, showing how plants vary in the wild and display adaptations. His two chapters on geographical distribution presented innovative ideas about historical plant geography, where he discussed the possibility of northern species pushing through the tropics at some former cool period, and the means of plant dispersal, including the transport of seeds by birds and currents. After the Origin of Species, Darwin studied orchids as a prime example of adaptation, in this regard as adaptations to ensure insect fertilization. Later he worked extensively on the cross-fertilization of plants, on insectivorous plants and the movements of plants. In all this work, he focused on the active physiological life of the organism. Darwin also performed plant experiments to investigate the origin of variations, polymorphism and hybrids; and worked on pollination, the evolution of sexes, and tropisms (plant hormones). His close friends Joseph Hooker (18171911), director of the Royal Botanic Gardens, Kew, and Asa Gray (1818

1888), Professor of Botany at Harvard University, were both converted to evolutionary theory and defended Darwins ideas in reviews and in their botanical writings. Gray and Hooker were among the rst to use natural selection for understanding plant relationships. Generally, however, there were few ghts over the evolution of plants in the same way as there were ghts over the possible connections between animals and humankind. Botanists probably found the idea of plant evolution far less repugnant than that of human evolution, and did not worry too much about the moral and theological implications of plant descent. Even so, there were a number of satirical references and cartoons poking fun at the transformation of turnips into humans. The morphology of plants was generally underrated until evolutionary theory gave it new legitimacy. The idea of the alternation of generations in animals helped Wilhelm Hofmeister (18241877) demonstrate the two phases in the life cycle of mosses and liverworts in 1851. Hofmeister was followed by Ladislav Celakowsky who in 1868 coined the terms gametophyte and sporophyte. Celakowsky saw the evolutionary signicance of Hofmeisters work, arguing from the phylogenetic perspective that sexual generation must be more primitive than asexual. Heated debate arose over the origin of owering plants (angiosperms) in the Cretaceous period. This also involved controversy over what should properly be considered primitive. Throughout the nineteenth century Englers arrangement was enshrined in most classication schemes until C. E. Bessey, in 1915, proposed that Ranunculus and its relatives came closest among living plants to primitive angiosperms. Frederick Bower (18551948) linked the migration of plants onto land to the appearance and subsequent elaboration of an asexual generation. In 1868 the rst edition of Julius von Sachs Lehrbuch der Botanik appeared which led the way in plant physiological investigations in Europe. Sachs impact was as great as Linnaeuss some hundred years before although in a very dierent eld. Every eminent plant physiologist was trained in Sachs laboratory in Wurzburg. In many ways the state of English botany before then resembled that of English physiology. Both disciplines were founded on a descriptive anatomical tradition; both imported the new laboratory work wholesale from the continent; and both found it hard to become separate from the medical curriculum. Experimental botany at rst failed to nd a secure home in the English universities, with the exception of John Burdon Sanderson at University College London and Sidney Vines in Cambridge. Bowers appointments in London, and then Glasgow in 1885 revolutionized the way plant physiology was regarded. Among other achievements, Sachs made highly signicant studies on plant hormones, growth mechanisms and tropisms, such as the eect of gravity or light on roots and shoots. Elsewhere, the emphasis of university botany shifted so much towards the physiological, for example in enzyme
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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 Benthams Genera Plantarum. Botany went on to become the prime area for genetic research, as in pure line experiments, and cytology, fertilization and cell division.

Further Reading
Allan MA (1977) Darwin and his Flowers. The Key to Natural Selection. London: Faber and Faber.

Andrews HN (1980) The Fossil Hunters: In Search of Ancient Plants. Ithaca, NY: Cornell University Press. Delaporte F (1982) Natures Second Kingdom. Explorations of Vegetality in the Eighteenth Century, Goldhammer A (transl.). Cambridge, MA: MIT Press. Frey KJ (ed.) (1994) Historical Perspectives in Plant Science. Ames: Iowa State University Press. Green JR (1909) A History of Botany, 1860 1900. Being a Continuation of Sachs History of Botany 1530 1860. Oxford: Clarendon Press. Morton AG (1981) History of Botanical Science. An Account of the Development of Botany from Ancient Times to the Present Day. London: Academic Press.

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