Origins and concept
Hairston, Smith and Slobodkin (1960) asked why the world is green and argued that plants are not eaten away because predators keep herbivores in check. The idea is known as the HSS or green world hypothesis, after the authors' initials. On the rocky shore of Washington State, Robert Paine (1966) observed that removing a predatory starfish allowed mussels to cover the rocks and crowd out other species, and from this he developed the concept of the keystone species. Paine (1980) used the term trophic cascade for such chains of effects. Oksanen et al. (1981) formalized a theory in which food chains lengthen and their regulation changes as primary productivity rises.
Classic examples
Estes and Palmisano (1974) showed that in the Aleutian Islands, islands with sea otters had few sea urchins and lush kelp forests, whereas on islands without otters urchins grazed the kelp down to urchin barrens. Carpenter, Kitchell and Hodgson (1985) reported that in lakes, piscivorous fish that reduce plankton-eating fish allow zooplankton to increase and algae to decline. Ripple and Beschta (2004) argued that after wolves were reintroduced to Yellowstone National Park, changes in elk foraging allowed willow and aspen to recover in some areas, emphasizing an ecology of fear in which predation risk itself reshapes ecosystems. The scale and causes of the Yellowstone case, however, remain debated among researchers.
Top-down versus bottom-up and significance
Hunter and Price (1992) argued that ecosystems are shaped by both top-down control by predators and bottom-up control by nutrients and primary production, with their relative importance varying by context. Polis and Strong (1996) noted that omnivory and complex food webs can dampen cascades. Pace et al. (1999) gathered evidence of trophic cascades across diverse ecosystems, and Shurin et al. (2002) found in a cross-ecosystem comparison that cascades tend to be stronger in aquatic than in terrestrial systems. The volume edited by Terborgh and Estes (2010) and Estes et al. (2011) warned that the loss of large predators is causing a trophic downgrading that simplifies ecosystems worldwide. The concept links population models such as the Lotka–Volterra equations with community theories such as the intermediate disturbance hypothesis.
FAQ
- What is a trophic cascade in simple terms?
- It is when an effect travels down a food chain: more top predators such as wolves mean fewer herbivores such as deer, which in turn lets trees and grasses increase.
- How do top-down and bottom-up control differ?
- Top-down control is when predators regulate the levels below them, while bottom-up control is when nutrients and plant production set the size of the levels above. In real ecosystems both forces operate together.
Sources reviewed: 2026-10-26
References
- Hairston, N. G., Smith, F. E., & Slobodkin, L. B. (1960). Community structure, population control, and competition. American Naturalist, 94(879), 421–425.
- Paine, R. T. (1966). Food web complexity and species diversity. American Naturalist, 100(910), 65–75.
- Paine, R. T. (1980). Food webs: Linkage, interaction strength and community infrastructure. Journal of Animal Ecology, 49(3), 667–685.
- Estes, J. A., & Palmisano, J. F. (1974). Sea otters: Their role in structuring nearshore communities. Science, 185(4156), 1058–1060.
- Carpenter, S. R., Kitchell, J. F., & Hodgson, J. R. (1985). Cascading trophic interactions and lake productivity. BioScience, 35(10), 634–639.
- Oksanen, L., Fretwell, S. D., Arruda, J., & Niemelä, P. (1981). Exploitation ecosystems in gradients of primary productivity. American Naturalist, 118(2), 240–261.
- Hunter, M. D., & Price, P. W. (1992). Playing chutes and ladders: Heterogeneity and the relative roles of bottom-up and top-down forces in natural communities. Ecology, 73(3), 724–732.
- Polis, G. A., & Strong, D. R. (1996). Food web complexity and community dynamics. American Naturalist, 147(5), 813–846.
- Pace, M. L., Cole, J. J., Carpenter, S. R., & Kitchell, J. F. (1999). Trophic cascades revealed in diverse ecosystems. Trends in Ecology & Evolution, 14(12), 483–488.
- Shurin, J. B., et al. (2002). A cross-ecosystem comparison of the strength of trophic cascades. Ecology Letters, 5(6), 785–791.
- Ripple, W. J., & Beschta, R. L. (2004). Wolves and the ecology of fear: Can predation risk structure ecosystems? BioScience, 54(8), 755–766.
- Terborgh, J., & Estes, J. A. (Eds.). (2010). Trophic Cascades: Predators, Prey, and the Changing Dynamics of Nature. Washington, DC: Island Press.
- Estes, J. A., et al. (2011). Trophic downgrading of planet Earth. Science, 333(6040), 301–306.
