ECOSYSTEM

the complex of living organisms, their physical environment, and all their interrelationships in a particular unit of space. The principles underlying the study of ecosystems are based on the view that all the elements of a life-supporting environment of any size, whether natural or man-made, are parts of an integral network in which each element interacts directly or indirectly with all others and affects the function of the whole. All ecosystems are contained within the largest of them, the ecosphere, which encompasses the entire physical Earth (geosphere) and all of its biological components (biosphere). An ecosystem can be categorized into its abiotic constituents, including minerals, climate, soil, water, sunlight, and all other nonliving elements, and its biotic constituents, consisting of all its living members. Linking these constituents together are two major forces: the flow of energy through the ecosystem, and the cycling of nutrients within the ecosystem. The fundamental source of energy in almost all ecosystems is radiant energy from the sun. The energy of sunlight is used by the ecosystem's autotrophic, or self-sustaining, organisms. Consisting largely of green vegetation, these organisms are capable of photosynthesisi.e., they can use the energy of sunlight to convert carbon dioxide and water into simple, energy-rich carbohydrates. The autotrophs use the energy stored within the simple carbohydrates to produce the more complex organic compounds, such as proteins, lipids, and starches, that maintain the organisms' life processes. The autotrophic segment of the ecosystem is commonly referred to as the producer level. Organic matter generated by autotrophs directly or indirectly sustains heterotrophic organisms. Heterotrophs are the consumers of the ecosystem; they cannot make their own food. They use, rearrange, and ultimately decompose the complex organic materials built up by the autotrophs. All animals and fungi are heterotrophs, as are most bacteria and many other microorganisms. Together, the autotrophs and heterotrophs form various trophic (feeding) levels in the ecosystem: the producer level, composed of those organisms that make their own food; the primary-consumer level, composed of those organisms that feed on producers; the secondary-consumer level, composed of those organisms that feed on primary consumers; and so on. The movement of organic matter and energy from the producer level through various consumer levels makes up a food chain. For example, a typical food chain in a grassland might be grass (producer) mouse (primary consumer) snake (secondary consumer) hawk (tertiary consumer). Actually, in many cases the food chains of the ecosystem overlap and interconnect, forming what ecologists call a food web. The final link in all food chains is made up of decomposers, those heterotrophs that break down dead organisms and organic wastes. A food chain in which the primary consumer feeds on living plants is called a grazing pathway; that in which the primary consumer feeds on dead plant matter is known as a detritus pathway. Both pathways are important in accounting for the energy budget of the ecosystem. As energy moves through the ecosystem, much of it is lost at each trophic level. For example, only about 10 percent of the energy stored in grass is incorporated into the body of a mouse that eats the grass. The remaining 90 percent is stored in compounds that cannot be broken down by the mouse or is lost as heat during the mouse's metabolic processes. Energy losses of similar magnitude occur at every level of the food chain; consequently, few food chains extend beyond five members (from producer through decomposer), because the energy available at higher trophic levels is too small to support further consumers. The flow of energy through the ecosystem drives the movement of nutrients within the ecosystem. Nutrients are chemical elements and compounds necessary to living organisms. Unlike energy, which is continuously lost from the ecosystem, nutrients are cycled through the ecosystem, oscillating between the biotic and abiotic components in what are called biogeochemical cycles. Major biogeochemical cycles include the water cycle, carbon cycle, oxygen cycle, nitrogen cycle, phosphorus cycle, sulfur cycle, and calcium cycle. Decomposers play a key role in many of these cycles, returning nutrients to the soil, water, or air, where they can again be used by the biotic constituents of the ecosystem. The orderly replacement of one ecosystem by another is a process known as ecosystem development, or ecological succession. Succession occurs when a sterile area, such as barren rock or a lava flow, is first colonized by living things or when an existing ecosystem is disrupted, as when a forest is destroyed by a fire. The succession of ecosystems generally occurs in two phases. The early, or growth, phase is characterized by ecosystems that have few species and short food chains. These ecosystems are relatively unstable but highly productive, in the sense that they build up organic matter faster than they break it down. The ecosystems of the later, or mature, phase are more complex, more diversified, and more stable. The final, or climax, ecosystem is characterized by a great diversity of species, complex food webs, and high stability. The major energy flow has shifted from production to maintenance. Human interference in the development of ecosystems is widespread. Farming, for example, is the deliberate maintenance of an immature ecosystemone that is highly productive but relatively unstable. Sound management of ecosystems for optimal food production should seek a compromise between the characteristics of young and mature ecosystems, and should consider factors that affect the interaction of natural cycles. Short-term production can be maximized by adding energy to the ecosystem in the form of cultivation and fertilization. Such efforts, however, can hinder efficient energy use in the long run by producing an imbalance of nutrients, an increase in pollutants, or a heightened susceptibility to plant diseases as a consequence of intensive inbreeding of crops. Although an awareness of the interdependence between human society and its environment was already prominent in ancient philosophy and religion, the formulation of the basic principles of systems ecology as a scientific discipline began in the late 19th century. During the second half of the 20th century, the study of ecosystems has become increasingly sophisticated and is now instrumental in the assessment and control of the effects of agricultural development and industrialization on the environment. On farms, for instance, it has shown that optimal long-term production of pasturage requires a moderate grazing schedule in order to ensure a steady renewal of the moisture and nutrient content of the soil and has emphasized the need for multiple-use strategies in the cultivation of arable lands. Systems ecology has been concerned with the consequences of accumulated insecticides and has provided a way of monitoring the climatic effects of atmospheric dust and carbon dioxide released by the burning of fossil fuels (e.g., coal, oil, and natural gas). It has helped to determine regional population capacities and has furthered the development of recycling techniques that may become essential in humanity's future interaction with the environment.