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Many theories have been formulated to explain the role of grazers such as zooplankton in controlling the amount of planktonic algae (phytoplankton) in lakes. The first theories of such grazer control were merely based on observations of negative correlations between algal and zooplankton numbers. A low number of algal cells in the presence of a high number of grazers suggested,but did not prove,that the grazers had removed most of the algae. The converse observation, of the absence of grazers in areas of high phytoplankton concentration, led Hardy to propose his principle of animal exclusion, which hypothesized that phytoplankton produced a repellent that excluded grazers from regions of high phytoplankton concentration. This was the first suggestion of algal defenses against grazing.

Perhaps the fact that many of these first studies considered only algae of a size that could be collected in a net (net phytoplankton), a practice that overlooked the smaller phytoplankton (nannoplankton) that we now know grazers are most likely to feed on, led to a de-emphasis of the role of grazers in subsequent research. Increasingly, as in the individual studies of Lund, Round, and Reynolds, researchers began to stress the importance of environmental factors such as temperature, light, and water movements in controlling algal numbers. These environmental factors were amenable to field monitoring and to simulation in the laboratory. Grazing was believed to have some effect on algal numbers, especially after phytoplankton growth rates declined at the end of bloom periods, but grazing was considered a minor component of models that predicted algal population dynamics.

The potential magnitude of grazing pressure on freshwater phytoplankton has only recently been determined empirically. Studies by Hargrave and Geen estimated natural community graz-ing rates by measuring feeding rates of individual zooplankton species in the laboratory and then computing community grazing rates for field conditions using the known population density of grazers. The high estimates of grazing pressure postulated by these researchers were not fully accepted, however, until the grazing rates of zooplankton were determined directly in the field, by means of new experimental techniques. Using a specially prepared feeding chamber, Haney was able to record zooplankton grazing rates in natural field conditions. In the periods of peak zooplankton abundance, that is, in the late spring and in the summer, Haney recorded maximum daily community grazing rates, for nutrient-poor lakes and bog lakes, respectively, of 6.6 percent and 114 percent of daily phytoplankton production. Cladocerans had higher grazing rates than copepods, usually accounting for 80 percent of the community grazing rate. These rates varied seasonally, reaching the lowest point in the winter and early spring.Haney's thorough research provides convincing field evidence that grazers can exert significant pressure on phytoplankton population.

The author most likely mentions Hardy's principle of animal exclusion in order to

give an example of one theory about the interaction of grazers and phytoplankton defend the first theory of algal defenses against grazing support the contention that phytoplankton numbers are controlled primarily by environmental factors demonstrate the superiority of laboratory studies of zooplankton feeding rates to other kinds of studies of such rates refute researchers who believed that low numbers of phytoplankton indicated the grazing effect of low numbers of zooplankton

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