ecology

ecology, study of the relationships between organisms and their environment. Some of the most pressing problems in human affairs—expanding populations, food scarcities, environmental pollution including global warming, extinctions of plant and animal species, and all the attendant sociological and political problems—are to a great degree ecological.

The word ecology was coined by the German zoologist Ernst Haeckel, who applied the term oekologie to the “relation of the animal both to its organic as well as its inorganic environment.” The word comes from the Greek oikos, meaning “household,” “home,” or “place to live.” Thus, ecology deals with the organism and its environment. The concept of environment includes both other organisms and physical surroundings. It involves relationships between individuals within a population and between individuals of different populations. These interactions between individuals, between populations, and between organisms and their environment form ecological systems, or ecosystems. Ecology has been defined variously as “the study of the interrelationships of organisms with their environment and each other,” as “the economy of nature,” and as “the biology of ecosystems.”

Historical background

Ecology had no firm beginnings. It evolved from the natural history of the ancient Greeks, particularly Theophrastus, a friend and associate of Aristotle. Theophrastus first described the interrelationships between organisms and between organisms and their nonliving environment. Later foundations for modern ecology were laid in the early work of plant and animal physiologists.

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In the early and mid-1900s two groups of botanists, one in Europe and the other in the United States, studied plant communities from two different points of view. The European botanists concerned themselves with the study of the composition, structure, and distribution of plant communities. The American botanists studied the development of plant communities, or succession (see community ecology: Ecological succession). Both plant and animal ecology developed separately until American biologists emphasized the interrelation of both plant and animal communities as a biotic whole.

During the same period, interest in population dynamics developed. The study of population dynamics received special impetus in the early 19th century, after the English economist Thomas Malthus called attention to the conflict between expanding populations and the capability of Earth to supply food. In the 1920s the American zoologist Raymond Pearl, the American chemist and statistician Alfred J. Lotka, and the Italian mathematician Vito Volterra developed mathematical foundations for the study of populations, and these studies led to experiments on the interaction of predators and prey, competitive relationships between species, and the regulation of populations. Investigations of the influence of behaviour on populations were stimulated by the recognition in 1920 of territoriality in nesting birds. Concepts of instinctive and aggressive behaviour were developed by the Austrian zoologist Konrad Lorenz and the Dutch-born British zoologist Nikolaas Tinbergen, and the role of social behaviour in the regulation of populations was explored by the British zoologist Vero Wynne-Edwards. (See population ecology.)

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Biology Bonanza

(Read Thomas Malthus’s 1824 Britannica essay on population.)

While some ecologists were studying the dynamics of communities and populations, others were concerned with energy budgets. In 1920 August Thienemann, a German freshwater biologist, introduced the concept of trophic, or feeding, levels (see trophic level), by which the energy of food is transferred through a series of organisms, from green plants (the producers) up to several levels of animals (the consumers). An English animal ecologist, Charles Elton (1927), further developed this approach with the concept of ecological niches and pyramids of numbers. In the 1930s, American freshwater biologists Edward Birge and Chancey Juday, in measuring the energy budgets of lakes, developed the idea of primary productivity, the rate at which food energy is generated, or fixed, by photosynthesis. In 1942 Raymond L. Lindeman of the United States developed the trophic-dynamic concept of ecology, which details the flow of energy through the ecosystem. Quantified field studies of energy flow through ecosystems were further developed by the brothers Eugene Odum and Howard Odum of the United States; similar early work on the cycling of nutrients was done by J.D. Ovington of England and Australia. (See community ecology: Trophic pyramids and the flow of energy; biosphere: The flow of energy and nutrient cycling.)

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The study of both energy flow and nutrient cycling was stimulated by the development of new materials and techniques—radioisotope tracers, microcalorimetry, computer science, and applied mathematics—that enabled ecologists to label, track, and measure the movement of particular nutrients and energy through ecosystems. These modern methods (see below Methods in ecology) encouraged a new stage in the development of ecology—systems ecology, which is concerned with the structure and function of ecosystems.