II. Read the following text paying attention to the highlighted words. Explain or interpret the contextual meaning of the underlined phrases

Ecology, also called bioecology, bionomics, or environmental biology, studies the relationships between organisms and their environment.

Long unfamiliar to the public, and relegated to a second-class status by many in the world of science, ecology emerged in the late 20th century as one of the most popular and most important aspects of biology. It has become painfully evident that some of the most pressing problems in the affairs of men — expanding populations, food scarcities, environmental pollution, and all the attendant sociological and political problems — are to a great degree ecological.

The word ecology was coined by a 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 word 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.”

In the early and mid-1900s two groups of botanists, one in Europe and the other in America, 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. 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 Thomas Malthus called attention to the conflict between expanding populations and the capability of the earth to supply food. R. Pearl (1920), A.J. Lotka (1925), and V. Volterra (1926) 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. Investigation of the influence of behaviour on populations was stimulated by the recognition in 1920 of territoriality in nesting birds. Concepts of instinctive and aggressive behaviour were developed by K. Lorenz and N. Tinbergen, and the role of social behaviour in the regulation of populations was explored by V.C. Wynne-Edwards.

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, 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, C.E. Elton (1927), further developed this approach with the concept of ecological niches and pyramids of numbers. Two American freshwater biologists, E. Birge and C. Juday, in the 1930s, in measuring the energy budgets of lakes, developed the idea of primary production, i.e., the rate at which food energy is generated, or fixed, by photosynthesis. Modern ecology came of age in 1942 with the development, by R.L. Lindeman of the United States, of 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 Eugene 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.

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

Until the late 20th century ecology lacked a strong conceptual base. Modern ecology, however, is now focussed on the concept of the ecosystem, a functional unit consisting of interacting organisms and all aspects of the environment in any specific area. It contains both the nonliving (abiotic) and living (biotic) components through which nutrients are cycled and energy flows. To accomplish this cycling and flow, ecosystems must possess a number of structured interrelationships between soil, water, and nutrients, on the one hand, and producers, consumers, and decomposers on the other. Ecosystems function by maintaining a flow of energy and a cycling of materials through a series of steps of eating and being eaten, of utilization and conversion, called the food chain. Ecosystems tend toward maturity, or stability, and in doing so they pass from a less complex to a more complex state. This directional change is called succession. Whenever an ecosystem is used, and that exploitation is maintained — as when a pond is kept clear of encroaching plants or a woodland is grazed by domestic cattle — the maturity of the ecosystem is effectively postponed. The major functional unit of the ecosystem is the population. It occupies a certain functional niche, related to its role in energy flow and nutrient cycling. Both the environment and the amount of energy fixation in any given ecosystem are limited. When a population reaches the limits imposed by the ecosystem, its numbers must stabilize or, failing this, decline from disease, starvation, strife, low reproduction, or other behavioural and physiological reactions. Changes and fluctuations in the environment represent selective pressure upon the population to which it must adjust. The ecosystem has historical aspects: the present is related to the past and the future to the present. Thus the ecosystem is the one concept that unifies plant and animal ecology, population dynamics, behaviour, and evolution.

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