Environmental Engineering Reference
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through the Gordian knot of ecosystem complexity, and reliably show how the system
actually reacts to some perturbation. It took a few decades for such whole-ecosystem
experiments to become a common and accepted tool. Perhaps because of the pervasive
influence of “The Lake as a Microcosm” and the clear boundaries to lakes, many of the
earliest whole-ecosystem experiments were performed on lakes ( Likens 1985; Carpenter
et al. 1995 ). For instance, models and small-scale experiments were unable to resolve a bit-
ter controversy in the 1960s about whether excessive phosphorus caused lakes to become
offensively eutrophic, but a whole-lake experiment was conclusive (see Figure 8.1 in
Chapter 8). Likewise, by adapting the small-watershed technique from hydrology in the
1960s, ecosystem scientists could quantify inputs and outputs of materials to and from ter-
restrial ecosystems and treat entire watersheds as experimental subjects ( Bormann and
Likens 1967; Likens et al. 1970 ). Perhaps more than any other tool, whole-ecosystem
experiments made Tansley's concept a practical subject of scientific study. Ecosystem
experiments are now an important tool for scientists to study subjects as varied as the
effects of toxins, food-web structures, disturbances, and limiting nutrients in all types of
ecosystems ( Table 1.1 ).
As a result of these advances, during the period from approximately 1935 to 1975, eco-
system science moved from being just an interesting concept to a central position in con-
temporary ecology. Ecosystem scientists, from the roots of the discipline to the present,
have worked to unravel the complexity of entire ecosystems of all sizes and forms, from a
water-filled cavity in a tree, to a small vernal pool, to a large lake, to a forested watershed,
to an entire city, to the total biosphere. The ecosystem concept provides a comprehensive
framework for study of the interactions among individuals, populations, and communities
and their abiotic environments, and for study of the change in these relationships both
temporally and spatially (adapted from Likens 1992 ).
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