Cause-Effect Relationships in Energy Flow, Trophic Structure, and Interspecific Interactions

Measurements of the efficiency of energy transfer between trophic levels are consis- tent with the hypothesis that it is trophic structure that controls the fraction of energy consumed at each trophic level, rather than energetics controlling trophic structure. Moreover, trophic structure is determined by competitive and predator-prey interactions. In freshwater pelagic communities, the collective efficiency of herbivorous plankton in consuming primary producers is up to 10 times as great as is the efficiency of forest herbivores in consuming their food. Conversely, forest predators are about three times as efficient in consuming herbivore produc- tion as are zooplanktivorous fish. The presence of an additional level, piscivorous fish, in pelagic communities accounts for the difference. In the aquatic system, herbivorous zooplankton are freed from predation by the effect of piscivorous fish on their predators; in the terrestrial system, green plants are freed from herbivory by predation on the herbivores. We explain the contrast between freshwater pelagic systems and forests and prairies as follows: Pelagic ecosystems have more trophic levels as a result of selection for small rapidly growing primary producers, which cannot hold space in the fluid medium, in contrast to large space-occupying producers in the terrestrial environment. Consumers in pelagic systems are more frequently gape limited in the size range of food they can ingest than are grasping consumers in terrestrial systems. This difference makes for two largely distinct levels of predators in pelagic communities. The energy within the living, nondetrital components is more finely divided between trophic levels in pelagic systems than in terrestrial systems. Ecological efficiencies do not determine trophic structure; rather, they are its product.