Specialized Sciences The 1800s

Early in the century, Alexander von Humboldt (Figure 10) became famous for his investigations and publications. He used inherited wealth to fund 5 years of exploration in Latin America, and then spent two decades in Paris, publishing his findings. 'Humboldtian science' involved correlations between plant associations (e.g., grasslands, rain forests, tundra) and environmental factors (e.g., temperature, precipitation, topography), which he carefully measured to understand the distribution and abundance of species of plants and animals. He was the effective founder of phytogeography, emphasizing distribution of vegetation rather than floristics.

In 1809, Lamarck (Figure 11) published his theory of evolution, arguing that species do not become extinct; they become different species. August de Candolle (Figure 12), who rejected Lamarck's theory, emphasized (1820) competition between species as important for determining distributions and extinctions. His writings

Figure 10 Alexander von Humboldt (1769-1859).

Jean Baptiste Lamarck
Figure 11 Jean Baptiste de Lamarck (1744-1829).
Jean Baptiste Lamarck 1809
Figure 12 August-Pyramus de Candolle (1778-1841).
Lamarck Figure
Figure 13 Hewett Cottrell Watson (1804-81).

were used by geologist Charles Lyell (1832), who believed in the stability of species and that fossils do represent extinct species.

Hewett C. Watson (Figure 13), founder of British phytogeography, was strongly influenced by Humboldt

Figure 14 Charles Robert Darwin (1809-82).

and Lamarck but not by de Candolle. He investigated the variability of British plant species in different parts of their ranges, and also compared the British and Azores Island floras.

Charles Darwin (Figure 14), who read widely, was already familiar with the argument for the importance of competition in nature before he read in 1837 Thomas R. Malthus' Essay on the Principle of Population (1798, 5th edn. 1825). Darwin's theory of evolution by natural selection focused attention on the struggle of organisms against biotic and abiotic environmental forces. In such a revolutionary book as On the Origin of Species (1859), one might expect Darwin to reject the balance of nature concept, since the struggle for existence can cause extinctions. However, his famous cats-mice-bees-clover story seemed to support the balance concept. It was Alfred Russel Wallace who, thinking of extinctions, asked ''Where is the balance?''

A Darwin disciple, Ernst Haeckel (Figure 15), in 1866 expressed the need for a new science, 'oecology'. In 1870, he explained, ''By oecology we mean the body of knowledge concerning the economy of nature - the investigation of the total relations of the animal both to its inorganic and to its organic environment.'' It took about three more decades for ecological sciences to begin to organize, but meanwhile, various ecological

Ernst Haeckel
Figure 15 Ernst Heinrich Haeckel (1834-1919).

ideas were elaborated. Darwin's theory of evolution included positive relations between individuals and between species, but the emphasis seemed to be on the negative relations of competition, predation, and parasitism. Several authors explored positive relations: Anton de Bary coined the word 'symbiosis' and wrote a book on it (1869); P.J. van Beneden defined mutalism in his book Les commensaux et les parasites (1875); Alfred Espinas described diverse forms of mutualism in Des societes animals (1878); and Peter Kropotkin wrote on both animals and humans in Mutual Aid (1902). Parasitology and the germ theory of disease were important developments in the late 1800s, but their relevance for ecology was little developed.

Edward Forbes (Figure 16) began studying marine animals in the 1830s and published many works concerning the distribution of species in relation to depth and other factors. He postulated the existence of an 'azotic' zone in the depths of the sea. J. J. Coste established a marine zoology laboratory at Concarneau, France, in 1859; more influential was Anton Dohrn's Stazione Zoologica, established at Naples in 1872. The latter influenced Spencer Fullerton Baird to obtain funds for a US Fish Commission Laboratory at Woods Hole, Massachusetts, in 1884. In 1888, the independent Woods Hole Oceanographic Institution opened nearby. Charles Wyville Thomson (Figure 17) was the first student of deep oceanic life, and his book, The Depths of the Sea

Figure 16 Edward Forbes, Jr. (1815-54).

Wyville Thomson Hms Porcupine
Figure 17 Charles Wyville Thomson (1830-82).
Stephen Alfred Forbes
Figure 18 Stephen Alfred Forbes (1844-1930).

(1872), helped him become head naturalist on the worldwide voyage of the HMS Challenger (1872-76), and later he directed publication of 50 large volumes of its scientific reports (1885-95). Impressive discoveries of this expedition inspired other countries to sponsor oceanographic voyages. Karl Mobius (German 1877, English 1883) studied oyster beds to assist in their management. He argued that an oyster bed is a ''community of living beings, a collection of species and a massing of individuals, which find here everything necessary for their growth and continuance.'' His study was influential for developing the concept of a biotic community.

American zoologist Stephen A. Forbes (Figure 18) wrote an influential essay, 'The Lake as a Microcosm' (1887), which may have been influenced by Mobius' essay, and which used the balance of nature concept to explain the apparent stability of species in a lake despite competition and predation. Meanwhile, since the 1860s, the Swiss zoologist Francois Alphonse Forel (Figure 19) devoted his research to the life in Lake Geneva and the environmental conditions. His studies culminated in a three-volume paradigm treatise on this research (18921904), which he named as 'limnology', and he published the first textbook on it in 1901.

Although Swiss botanist Carl Schroter first introduced the terms 'autecology' and 'synecology' in 1910, those subjects already existed by the 1890s. A strong German tradition in plant physiology and phytogeography led to

Fran Ois Alphonse Forel
Figure 19 Frangois Alphonse Forel (1841-1912).

the development of plant autecology, and when a botanical congress in Madison, Wisconsin, in 1893 adopted the term 'ecology', 'autecology' was meant. However, the first plant ecology treatise, by Eugenius Warming (Figure 20), Plantesamfund (1895), was on synecology, based on his lectures in the first course ever taught in ecology, at the University of Copenhagen. Also in 1895 S. A. Forbes claimed that ''economic entomology is simply applied ecology.''

The 1900s Specializations

Four specializations of ecology that emerged during the late 1800s - plant ecology, animal ecology, limnology, and marine ecology - persisted throughout the 1900s, but other specializations also arose, most notably population ecology and ecosystem ecology (Figure 21). Population ecology focused primarily on animals, but both plant and animal ecologists developed ecosystem ecology.

American plant ecologists Frederic E. Clements and Henry Chandler Cowles (Figures 22 and 23) led the investigation of plant communities. Clements wrote the first ecology textbook in English (1905), which was used in the US and in the British Empire. Clements believed

Arthur Tansley Ecology

Figure 20 Johannes Eugenius Warming (1841-1924). Courtesy of Hunt Institute for Botanical Documentation.

plant communities were united into a 'superorganism', but in 1917 H. A. Gleason questioned the existence of biotic communities, and in 1926 he developed 'The Individualistic Concept of the Plant Association'. Arthur G. Tansley (Figure 24) at Oxford University challenged the concept of a community being a superorganism in 'The Use and Abuse of Vegetational Concepts and Terms' in 1935. Gleason's perspective was adopted in the later 1940s by John T. Curtis (Figure 25) in his school of plant ecology at the University of Wisconsin, where they collected quantitative data for gradient analysis to document a 'continuum' of plant species distributions.

In limnology there was a strong interest in classifying lakes according to their biological productivity and in measuring their physical and chemical qualities. American leadership was by Edward A. Birge and Chauncey Juday at the University of Wisconsin; for many Wisconsin lakes, they studied temperature stratification, light penetration, dissolved minerals, and hydrogen ion concentration. Arthur D. Hasler (Figure 26), who succeeded them, emphasized experimental limnology and fish ecology. G. Evelyn Hutchinson (Figure 27) of Yale University published an encyclopedic Treatise on Limnology (1957-67). In Europe, the leading limnologist was August Thienemann (Figure 28), at the Hydrobiologische Anstalt at Lake Plon and at the University of Kiel. He studied the physical qualities of north German lakes and their invertebrate fauna, and in 1928-29, to make comparisons, he led an expedition to Indonesia to study life in tropical lakes.

The first attempts to organize animal ecology was by Charles C. Adams, who taught animal ecology at the University of Illinois and published a Guide to the Study of Animal Ecology (1913), and by Victor E. Shelford (Figure 29), who taught animal ecology at the University of Chicago and published Animal Communities in Temperate America as Illustrated in the Chicago Region (1913). A more sweeping synthesis was by Charles Elton (Figure 30) of Oxford University, whose Animal Ecology (1927) explained the dynamics of food chains and transformed Joseph Grinnell's descriptive concept (1917) of the niche of California thrashers into a functional concept. In 1939, Clements and Shelford collaborated on a textbook, Bio-Ecology, which sought to bridge the gaps between their specializations. It helped, though the specializations persisted. In 1949, five Chicago animal ecologists - Warder C. Allee, Alfred E. Emerson, Orlando Park, Thomas Park, and Karl P. Schmidt -collaborated on the most detailed and comprehensive synthesis ever written (including two historical chapters), Principles ofAnimal Ecology, which emphasized cooperation in nature.

Marine ecology focused on estimating the abundance of plankton, at the bottom of the food chain, and fish at the top, seeking in both cases causes of fluctuation in abundance. Advances in commercial fishing technology after the Second World War created an industry that could deplete oceans and great lakes of their resources, and ecologists and fishery biologists were needed to explain the level of harvesting that can be sustained. The same is true for shellfish, which must also be protected from pollution. Hasler's study of homing instinct in salmon facilitated development of 'salmon farming'.

Tansley named and defined the 'ecosystem' concept in 1935 but did not lay its foundation. (Instead, he completed his large monograph, The British Islands and Their Vegetation, in 1939.) Raymond L. Lindeman, a Midwestern postdoctoral student under Hutchison, developed the ecosystem concept in the most important ecological article ever published, 'The Trophic-Dynamic Aspect of Ecology' (1942). It includes both original research and a synthesis of previous literature. Lindeman died, but Hutchinson got it published. After the Second World War, the brothers Eugene and Howard Thomas Odum (Figure 31) led in studies on the productivity of ecosystems. Eugene Odum also published a very

Ecologie

Margalef Von neumann

Climatologie Écologie Cycles Écologie des Écologie globale biogeochimiques populations génétique

Ecologie Phytosociologie Biogéographie Ecologie du systémique paysage planétaire Lovelock May ri_ ,,

Margulis [Boudyko

Écologie moderne

Naissance de l'écologie 1900

1850 Racines de l'écologie

L' économie de la nature planétaire Lovelock May

Margalef Von neumann

Mathématique du chaos

Sukhatchev Pédologie

Fourier

Agrochimie Dumas Boussingault Liebig

Fourier

Priestley Physiocrates

Priestley Physiocrates

Mathématique du chaos

Gause

Ostwald oukouchaev linsky

Climatologie Écologie Cycles Écologie des Écologie globale biogeochimiques populations génétique

Howard Margulis

Sukhatchev Pédologie

Approche populationnelle

Approche botanique

Gause

Kasharov

Volterra Lotka Vernadsky Anapm~\i

Ross

Physique Marchal mathématique

Ostwald

Arrhenius Révolution Pédologie Bo„zmann Génétiqi pastorienne russe Bonzmann oukouchaev

Thermodynamique Haeckel linsky pastorienne

Doukouchaev Pasteur Thermodynamique

Podolinsky

Agrochimie Dumas Boussingault Liebig

Approche physico-chimique

Approche populationnelle

Insectologie Réaumur

Approche botanique

Figure 21 Some roots and branches of the tree of ecological knowledge (Deléage 1992). It includes names of some ecologists not mentioned in this article. Courtesy of Éditions la Découverte.

Tansley Ecologia
Figure 22 Frederic Édward Clements (1874-1945). Courtesy of Hunt Institute for Botanical Documentation.
Henry Chandler Cowles
Figure 23 Henry Chandler Cowles (1869-1939).

1950

1925

1800

Georges Ostwalt
Figure 24 Arthur George Tansley (1871-1955). Courtesy of Hunt Institute for Botanical Documentation.

popular textbook, Fundamentals of Ecology (1953, 3d edn. 1971) that publicized ecosystem concepts. His ecology school at the University ofGeorgia studied the ecosystem within the vast US Atomic Energy Commission's Savannah River Installation Area. H. T. Odum pioneered ecosystem modeling, and today modeling is fully integrated into ecology and applied ecology due to important contributions from Denmark, the Netherlands, Sweden, Switzerland, and the US. Recent integrated ecosystem theory is indebted to H. T. Odum (maximum power and energy), R. Ulanowicz (ascendency), B. C. Patten (network theory), and thermodynamics (S. E. J0rgensen). R. O'Neill and Tim Allen developed hierarchy theory.

Population studies focused primarily on causes of fluctuations in abundance and on developing accurate mathematical models to describe and predict these fluctuations. The studies were mainly on mammals, birds, fish, and insects. Charles Elton led members of his Bureau of Animal Population (1932-67) into field work and theoretical debate at Oxford University. He discredited the notion of a balance of nature and replaced it with the idea of constant change. Aspects of ethology, the study of animal behavior, are relevant to animal ecology. The three founders of ethology in the 1930s (Konrad Lorenz, Niko Tinbergen, and Karl von Frisch) received a Nobel Prize in 1973.

The Russian geochemist Vladimir Ivanovich Vernadsky (Figure 32) studied organisms as the source of atmospheric gases and founded the science of biogeochemistry, explained in La biosphere (Russian 1926, French 1929, English 1997). Biogeochemistry has remained a dominant

Thomas Hunt 1800s
Figure 25 John Thomas Curtis (1913-61). Courtesy of Hunt Institute for Botanical Documentation.

aspect of Soviet/Russian ecology. A similar (but more radical) idea to Vernadsky's is James Lovelock's Gaia theory (1972), which postulates that the Earth is a superorganism that regulates its life forms to maintain an environment that supports life. Gaia theory has attracted popular support and continues to intrigue some Earth scientists, as seen in Scientists Debate Gaia (2004).

Ramon Margalef became the leading ecologist of Spain and the Spanish-speaking world with his 950-page Ecologia (1974, 4th edn. 1986), though he also published papers and Perspectives in Ecological Theory (1968) in English. Michael H. Graham, Paul K. Dayton, and Robert T. Paine have debated in Ecology (June 2002) whether ecological concepts are advanced according to a Popperian or Kuhnian model ofscience or by evolution of a paradigm - showing that a definitive answer probably requires a book-length study.

Mod Word Proc VerbalGeorge Evelyn Hutchinson

Figure 27 George Evelyn Hutchinson (1903-91). Courtesy of

Yale Peabody Museum of Natural History. Figure 28 August Friedrich Thienemann (1882-1960).

Figure 27 George Evelyn Hutchinson (1903-91). Courtesy of

Yale Peabody Museum of Natural History. Figure 28 August Friedrich Thienemann (1882-1960).

Societies and journals

The founding of societies and journals is an important indicator of scientific progress. The International Council for the Exploration of the Sea, founded in 1902 with headquarters in Copenhagen, pioneered in relating oceanography to fisheries. Its publications were

Journal du Conseil and Rapports et Proces Verbaux. The first ecological society arose in Britain. Tansley and William Smith had formed the British Vegetation Committee in 1904, which was very active but had difficulty finding funds to publish its memoirs and maps. They decided that an ecological society would be more successful in

Victor Ernest Shelford 1877 1968
Figure 29 Victor Ernest Shelford (1877-1968).
Betty Jean Craige
Figure 30 Charles Elton (1900-91).
Betty Jean Craige
Figure 31 Gene Elden Likens (1935-), Howard Thomas Odum (1924-2002), Eugene Plesants Odum (1913-2002), and William Eugene Odum (1942-1991). Courtesy of Betty Jean Craige and University of Georgia Press.
Betty Jean Craige
Figure 32 Vladimir Ivanovich Vernadsky (1863-1945).

attracting funds. The British Ecological Society held its first meeting on 12 April 1913, and the first issue of its Journal of Ecology was printed for that meeting. Tansley edited it, beginning from 1916. British animal ecology developed more slowly, and only in 1932 did the British Ecological Society begin publishing its Journal ofAnimal Ecology, edited by Elton. The Ecological Society of

America (encompassing USA and Canada) was founded in 1915; in 1917, it began publishing its Bulletin and in 1920 its main journal, Ecology. In 1922, Thienemann and Einar Naumann, a phytoplankton specialist at the University of Lund, founded the Internationale Vereinigung fuir Limnologie (later called Societas Internationalis Limnologiae), which meets biannually and publishes its proceedings. In 1936, Americans and Canadians founded the Limnological Society of America, which in 1948 became the American Society of Limnology and Oceanography. It began publishing Limnology and Oceanography in 1956. Since ecological papers were published in botanical, zoological, and general science journals, not all countries formed ecological societies, and some countries did so only later. Germany's Gesellschaft fuir Okologie was founded in 1970 and Italy's Societa Italiana di Ecologia in 1976. More than a dozen specialized ecological journals in English have begun since 1960.

The development of ecosystem models by H. T. Odum and his associates encouraged Sven Erik J0rgensen to lay the foundation for the journal Ecological Modeling in 1975. Volume 1 had 320 pages; current volumes contain some 4000 pages per year. This increase is illustrative of the explosion in all aspects of applied ecology since the last decades of the 1900s. Subjects initially published in Ecological Modeling soon acquired even more specialized journals: Ecological Economics (1988), Ecological Engineering (1992), Ecosystem Health (2001), Ecological Complexity (2004), Ecological Informatics (2006), and others.

Institutions

Ecology has flourished in both research institutions and university departments. The World Directory of Hydrobiological and Fisheries Institutions which Robert W. Hiatt compiled in 1963 provides information on hundreds of institutions, and their numbers have increased considerably since then. Recent lists of aquatic and terrestrial institutions are in Europa Publications' annuals, entitled The World of Learning. Here are just two American examples: the Marine Biological Association of San Diego established its laboratory in 1903, and it became the Scripps Institution of Oceanography in 1925; the Carnegie Institution of Washington in 1903 established the Carnegie Desert Botanical Laboratory outside Tucson; its funding declined during the Depression and in 1940 it was given to the US Forest Service, which sold it in 1960 to the University of Arizona. There are histories of several ecological schools: the plant ecology schools at the Universities of Nebraska and Chicago (Tobey); the Chicago animal ecology school (Mitman); each of the three schools at Oxford University: Tansley's plant ecology school (Anker), Elton's Population Bureau (Crowcroft), and Tinbergen's ethology school (Kruuk, Thorpe); the schools of plant ecology (Fraelish et al.) and limnology

(Becker and Egerton) at the University of Wisconsin-Madison; and Eugene Odum's ecosystem school at the University of Georgia (Barrett and Barrett; Craige).

Environmentalism

After World War II, both conservationists and ecolo-gists became concerned with negative aspects of the rapid human transformation of the environment. This was an opportunity for ecologists to advise on how to minimize environmental damage and preserve natural areas. This opportunity was first realized in the United Kingdom with the establishment of the Nature Conservancy in 1949. It both preserves natural areas and provides for ecological research in them. In the United States, the triple threats of atomic radiation, phosphate detergents, and dichlorodiphenyltrichlor-oethane (DDT) insecticide transformed pre-war conservationists into post-war environmentalists. Barry Commoner led the fight against atomic radiation and phosphate detergents, and Rachel Carson (Figure 33) led the fight against DDT. The outcome was passage of the National Environmental Policy Act (NEPA, 1969), requiring environmental impact assessments before

Rachel Carson 1970s
Figure 33 Rachel Louise Carson (1907-64).

major projects modify natural areas, and presidential establishment of the US Environmental Protection Agency (EPA, 1970), which monitors compliance with environmental laws. Both of these require expert advice from ecologists. Other countries have more or less followed the British and American examples.

The success of research conducted during the International Geophysical Year (1957-58) inspired three prominent scientists in 1959 to begin planning the International Biological Program, organized by the International Union of Biological Sciences. Barton Worthington (Figure 34), head of Britain's Nature Conservancy, became its scientific head. Nationally supported IBP research lasted from 1964 to 1974, and some projects continued beyond 1974. The projects were mainly ecological. In the United States, significant new funding enabled ecology to evolve into 'big science'. George M. Van Dyne and others used a method of monitoring ecosystems with computers called 'systems ecology'. Without new funds, the USSR contributed about as much IBP research and publications as did the USA. Worldwide, IBP held over 200 meetings, and its

Edgar Barton Worthington
Figure 34 Edgar Barton Worthington (1905-2001).

research cost over $40 million annually. The resulting publications were numerous and generally of high caliber.

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