History of Plant Ecology

Malcolm Nicolson, University of Glasgow, Glasgow, UK

Plant ecology is the study of the collective phenomena of plants. It dates from the nineteenth century, having its origins in Humboldtian plant geography and in Darwinian physiological anatomy. Much important work was done in the USA and Britain, both before and shortly after World War II. The study of plant communities is now a major aspect of ecosystem ecology.

Introductory article
Article Contents
. Introduction . Humboldtian Plant Geography . Physiological Anatomy . Plant Ecology in America and Britain . New Ecology

Introduction
The use of the term plant ecology, as the accepted name of a distinctive form of scientic inquiry, dates from the 1890s. But the history of the discipline extends considerably further back into the nineteenth century. Plant ecology has traditionally had two facets the study of how, and why, dierent plant species grow together in communities and the study of the relations of individual plant species to their environment. The former branch, known as synecology or vegetation science, is both the more dominant and the older partner, having its historical roots in the plant geography of Alexander von Humboldt. The latter branch, known as autoecology or physiological plant ecology, derives substantially from the impact of Darwinism upon mid-nineteenth-century plant anatomy and physiology.

example, an association in which one of the species of the oak, Quercus, was the most numerous tree, was termed a Quercetum. August Grisebach further developed Humboldts system of classication by growth forms and coined the term formation for a type of vegetation characterized solely by physiognomic criteria. In 1872, Grisebach published his great book, Die vegetation der Erde (The Vegetation of the Earth), a comprehensive treatise of physiognomic plant geography. By the 1880s, other plant geographers, in France, Switzerland and Scandinavia, had developed alternative systems of vegetational classication based upon more oristic criteria.

Physiological Anatomy
Following the appearance of Darwins On the Origin of Species in 1859, a number of plant physiologists, notably Simon Schwender at the University of Berlin, rebelled against the strict division between morphology and physiology which characterized German botany at that time and developed a programme of research in which a systematic attempt was made to correlate morphological structure with adaptive function, in the context of the life of the plant in the wild. Upon reading Die Vegetation der Erde, Schwender realized that his physiological anatomy, as it was termed, could potentially be applied to the investigation of the geographical distribution of plants and vegetation types. The challenge was to elucidate, from a physiological point of view, how particular plants were adapted to particular habitats. A similar research programme was developed simultaneously by Anton Schimper, a morphologist by training. German botanists were also inspired to elucidate the relationship between plants and the environment by their experience of exotic tropical vegetation. In the last decade of the nineteenth century, two major books were published Panzengeographie auf physiologischer Grundlage by Schimper and Lehrbuch der okologishen Panzen-geographie by Eugen Warming. Both books drew jointly upon Humboldtian plant geography and physiological anatomy to provide a comprehensive
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Humboldtian Plant Geography

Up until the end of the eighteenth century, botanists concentrated their eorts on nding individual plants and classifying the specimens into species and higher taxa. In 1790, however, Alexander von Humboldt set out a programme for a new form of plant geography which traces the connections and relations by which all plants are bound together among themselves, designates in what lands they are found, in what atmospheric conditions they live .... Humboldt drew attention to the fact that plants tended to grow together in recognizable and recurring groups and was the rst to use the term association, albeit informally, to describe these types of vegetation. He developed a non-oristic classication of vegetation types, based upon the growth form of the constituent plants. Using a number of measuring instruments, Humboldt sought to correlate the geographical distribution of vegetation with a wide variety of environmental variables. Humboldts eld work was done mainly in South America but later investigators, Anton Kerner for example, employed and rened his methods in the investigation of European vegetation. In 1822, Joachim Schouw devised the -etum nomenclature, whereby, for

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History of Plant Ecology

outline of what was now called plant ecology a programme of research into vegetation, its character, species composition and distribution; and into the relation of its constituent species to environmental factors. Also in the same decade, Oscar Drude, a student of Grisebach, produced his classic study, Deutchsland Panzengeographie. Schimper and Warmings books were quickly translated into English; the former as Plant Geography upon a Physiological Basis (1903), the latter as Oecology of Plants: An introduction to the study of plant communities (1909).

Plant Ecology in America and Britain

The inuence of these texts in English-speaking countries was immense. Field-orientated botanists, trained in oristics but aware of the decreasing fashionability of pure natural history, found ecological plant geography to be a fruitful expression of their skills and interests. In America rst Conway MacMillan, in Minnesota, and then Henry Cowles, in Chicago, were inspired by Warmings text to set out to classify their local vegetation along similar lines. Cowles, moreover, cleverly meshed the European systems of vegetational classication with the theory of geomorphological base-levelling which had been developed by American geologists such as his teacher, Rollin Salisbury. Warming had based his classications upon specic habitat factors such as the water content of the soil. For example, he recognized xerophytic, hydrophytic and mesophytic plant communities, associated respectively with dry, wet and moist soils the most developed form of vegetation being found in the intermediate conditions. But Cowles argued that soil conditions were themselves dependent upon a more fundamental environmental variable, namely topography. Soil water content was chiey determined by the slope and elevation of the terrain. Thus, as the continental landmass of America was gradually eroding toward the condition of a peneplain, so its vegetation was becoming more mesophytic in character, trending toward the most mesophytic vegetation type possible under the regional climate. This form of vegetation, deciduous woodland over much of temperate North America, was dubbed the climatic climax. The phenomena of succession, how types of vegetation replaced one another over time, had long been studied by botanists. Cowless theory, building upon this earlier work, conveyed a powerful sense both of the ubiquity of vegetational change and of an underlying predictable regularity within that change. It seemed to uncover the laws of vegetational development. And, perhaps most excitingly of all, it promised a privileged insight into the history of vegetation for:In many cases ... there is a horizontal order of succession at the present time that resembles the vertical [i.e. the temporal] succession of
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which we now have only the topmost member. [HC Cowles, 1901]. Thus by, for example, walking back from a sandy shore one could see, laid out in bands parallel with the waters edge, the development stages of the deciduous forest which was now established upon the stabilized and humuscovered mature sand dunes, furthest from the water. Cowles called his new approach physiographic ecology. His methods were rapidly applied throughout North America by his students, notably by W. S. Cooper at Glacier Bay in Alaska. Cooper became Professor of Ecology at the University of Minnesota, Minneapolis, and was one of the most inuential teachers of the subject in the United States. European examplars were also quickly adopted in the University of Nebraska, where Charles Bessey was Professor of Botany. Bessey was the leading American exponent of the so-called New Botany, the Germaninspired shift in emphasis from oristics and taxomony as the dominant modes of botanical instruction and inquiry towards the study of plants by means of microscopy and experimentation. The University of Nebraska was a LandGrant college and, in keeping with its institutional ethos, Bessey encouraged his students to pursue research topics that were both scientically worthwhile and economically useful. In particular, he was interested in the application of ecological knowledge to the practical problems of farming and ranching on the Great Plains. Besseys students, Roscoe Pound and Frederic Clements, adopted Drudes work as the model for their pioneering study, The Phytogeography of Nebraska. In 1905 Clements produced the textbook Research Methods in Ecology, which described the quantitative analysis of vegetation by the quadrat and transect techniques, which he and Pound had devised, and by the use of recording instruments in the eld. Clements research programme was developed further in an impressive series of books and articles, which culminated in 1916 with the publication of Plant Succession An Analysis of the Development of Vegetation. In Clements hands, academic plant ecology developed signicant links with agronomy, land management, horticulture and forestry. He served as a consultant to the Soil Conservation Service and the Great Plains Drouth Committee. In Britain, F. W. Oliver, Professor of Botany at University College London, translated Kerners work and developed the eld study of coastal plant communities. In 1899, his student and assistant, Arthur Tansley, began to teach a course based on Warmings Lehrbuch. In 1904, Tansley founded the British Vegetation Committee, under whose auspices the valuable guide book Types of British Vegetation was published in 1911. The Vegetation Committee formed the nucleus around which was founded, in 1913, the British Ecological Society the American equivalent dates from two years later. Tansley published the denitive The British Islands and their Vegetation in

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History of Plant Ecology

1939 and was the founding chairman of the Nature Conservancy Council. Throughout the late nineteenth and early twentieth centuries, nearly all vegetation research was predicated upon the conception that plants grow together in denite, ordered, repeating communities. These natural units of vegetation, characterized either by essentially constant physiognomy or essentially constant oristic composition, were held to constitute the proper subject matter for ecological study in the same way as plant species were the proper object of study for taxonomic botanists. Much European vegetation science, especially in southern Europe and Scandinavia, was based around the identication and classication of such units. In America, the leading proponent of the community unit theory was Frederic Clements, who claimed to discern in the formation an entity of organic, indeed super-organismic, integration. Henry Allan Gleason, however, argued that the plant community, while a valid object of study, was not an organism, scarcely even a vegetational unit but was merely the product of the co-incidence of plants able to live in the same location.

New Ecology
After World War II, the character of biological theorizing was transformed by advances in population genetics and by the development of the Neo-Darwinian synthesis. Clementsian ecology, speculative and with a pronounced Lamarckian character, became markedly old-fashioned. Detailed quantitative studies of the vegetation of Wisconsin, by John Curtis and his colleagues, and of the Great Smoky Mountains, by Robert Whittaker, vindicated Gleasons individualistic hypothesis, albeit now re-expressed with a more adaptive, Darwinian emphasis. Similar conclusions were arrived at by British botanists working in the tropical rainforest. These researches formed the departure point for the mathematically complex ordination and classication studies which were the cutting edge of vegetation science in the 1960s and early 1970s.

By the 1960s, however, plant ecology was, to a signicant extent, losing its distinctive disciplinary identity as a more inclusive science of general ecology emerged. In 1935, Tansley, partly as a corrective to what he regarded as the excesses of Clements super-organism theory, coined the term ecosystem. The new term conveyed the suggestion that plants, animals, fungi, bacteria and associated inanimate material should be studied together as components of an interrelated functioning entity. By the 1960s, ecologists had substantially taken up the challenge of investigating natural environments along these lines. Novel modes of investigation, such as the study of the ow of nutrients and energy, distinguished the New Ecology. Research techniques and equipment were increasingly borrowed from physics and chemistry, larger teams of workers were employed, and the use of computers for the manipulation of data and the modelling of processes became routine. Nevertheless, the concepts of succession and climax, as developed by plant ecologists, remained central to much ecosystem theory, as the work of Herbert Bormann and Gene Likens at Hubbard Brook Experimental Forest, New Hampshire exemplies. Meanwhile applied plant ecology continues to play a formative role in land management, wildlife conservation, forestry and related elds.

Further Reading
Cittadino E (1990) Nature as the Laboratory: Darwinian Plant Ecology in the German Empire 1800 1900. Cambridge: Cambridge University Press. Cowles HC (1901) The physiographic ecology of Chicago and vicinity; a study of the origin, development and classication of plant societies. Botanical Gazette 31: 73. Hagen JB (1992) An Entangled Bank: The Origins of Ecosystem Ecology. New Brunswick: Rutgers University Press. McIntosh RP (1985) The Background of Ecology: Concept and Theory. Cambridge: Cambridge University Press. Nicolson M (1996) Humboldtian plant geography after Humboldt: The link to ecology. British Journal for the History of Science 29: 289310. Whittaker RH (1975) Ordination and Classication of Communities. The Hague: Junk. Worster D (1985) Natures Economy: A History of Ecological Ideas. Cambridge: Cambridge University Press.

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