Trophic cascade

Trophic cascades occur when predators in a food chain suppress the abundance of their prey, thereby releasing the next lower trophic level from predation (or herbivory if the intermediate trophic level is an herbivore). For example, if the abundance of large piscivorous fish is increased in a lake, the abundance of their prey, zooplanktivorous fish, should decrease, large zooplankton abundance should increase, and phytoplankton biomass should decrease. This theory has stimulated new research in many areas of ecology. Trophic cascades may also be important for understanding the effects of removing top predators from food webs, as humans have done in many places through hunting and fishing activities.

Origins and theory

Nelson Hairston, Frederick E. Smith and Lawrence B. Slobodkin are generally credited with originating the concept of a trophic cascade, although they did not use the term. Hairston, Smith and Slobodkin argued that predators reduce the abundance of herbivores, allowing plants to flourish [Hairston NG, Smith FE, Slobodkin LB (1960) Community structure, population control and competition. American Naturalist 94:421-425] . This is often referred to as the green world hypothesis. The green world hypothesis is credited with bringing attention to the role of top-down forces (eg predation) and indirect effects in shaping ecological communities. The prevailing view of communities prior to Hairston, Smith and Slobodkin was trophodynamics, which attempted to explain the structure of communities using only bottom-up forces (eg resource limitation). Smith may have been inspired by the experiments of a Czech ecologist, Hrbáček, whom he met on a United States State Department cultural exchange. Hrbáček had shown that fish in artificial ponds reduced the abundance of zooplankton, leading to an increase in the abundance of phytoplankton [Hrbáček J, Dvořakova M, Kořínek V, Procházkóva L (1961) Demonstration of the effect of the fish stock on the species composition of zooplankton and the intensity of metabolism of the whole plankton association. Verh. Internat. Verein. Limnol. 14: 192-195] .

Hairston, Smith and Slobodkin argued that the ecological communities acted as food chains with three trophic levels of mud. Subsequent models expanded the argument to food chains with more than or fewer than three trophic levels [Oksanen L, Fretwell SD, Arruda J, Niemala P (1981) Exploitation ecosystems in gradients of primary productivity. American Naturalist 118:240-261] . Lauri Oksanen argued that the top trophic level in a food chain increases the abundance of producers in food chains with an odd number of trophic levels (such as in Hairston, Smith and Slobodkin's three trophic level model), but decreases the abundance of the producers in food chains with an even number of trophic levels. Additionally, he argued that the number of trophic levels in a food chain increases as the productivity of the ecosystem increases.

Criticisms

Although the existence of trophic cascades is not controversial, ecologists have long debated how ubiquitous they are. Hairston, Smith and Slobodkin argued that terrestrial ecosystems, as a rule, behave as a three trophic level trophic cascade, which provoked immediate controversy. Some of the criticisms, both of Hairston, Smith and Slobodkin's model and of Oksanen's later model, were:
#Opponents of pointed out that plants possess numerous defenses against herbivory, and that these defenses also contribute to reducing the impact of herbivores on plant populations Murdoch WM (1966) Community structure, population control, and competition -- a critique. American Naturalist 100:219-226] .
#Additionally, herbivore populations may be limited by factors other than food or predation, such as nesting sites or available territory.
#For trophic cascades to be ubiquitous, communities must generally act as food chains, with discrete trophic levels. Most communities, however, have complex food webs. In real food webs consumers often feed at multiple trophic levels (omnivory), organisms often change their diet as they grow larger, cannibalism occurs, and consumers are subsidized by inputs of resources from outside the local community, all of which blur the distinctions between trophic levels [Polis GA, Strong DR (1996) Food web complexity and community dynamics. American Naturalist 147: 813-846]

Classic examples

Although Hairston, Smith and Slobodkin formulated their argument in terms of terrestrial food chains, the earliest empirical demonstrations of trophic cascades came from marine and, especially, aquatic ecosystems. Some of the most famous examples are:
# In North American lakes, piscivorous fish can dramatically reduce populations of zooplanktivorous fish, zooplanktivorous fish can dramatically alter freshwater zooplankton communities, and zooplankton grazing can in turn have large impacts on phytoplankton communities. Removal of piscivorous fish can change lake water from clear to green by allowing phytoplankton to flourish [Carpenter SR, Kitchell JF, Hodgson JR (1985) Cascading trophic interactions and lake productivity. Bioscience 35:634-639] .
# In the Eel River, in Northern California, fish (steelhead and roach) consume fish larvae and predatory insects. These smaller predators prey on midge larvae, which feed on algae. Removal of the larger fish increases the abundance of algae [Power ME (1990) Effects of fish in river food webs. Science 250: 811-814] .
# In Pacific kelp forests, sea otters feed on sea urchins. In areas where sea otters have been hunted to extinction, sea urchins increase in abundance and decimate kelp [Estes JA, Palmisano JF (1974) Sea otters: their role in structuring nearshore communities. Science 185: 1058-1060]

Terrestrial trophic cascades

The fact that the earliest documented trophic cascades all occurred in lakes and streams lead Donald Strong to speculate that fundamental differences between aquatic and terrestrial food webs made trophic cascades primarily an aquatic phenomenon [Strong DR (1992) Are trophic cascades all wet? Differentiation and donor-control in speciose ecosystems. Ecology 73:747-754] . Strong argued that trophic cascades were restricted to communities with relatively low species diversity, in which a small number of species could have overwhelming influence and the food web could operate as a linear food chain. Additionally, well documented trophic cascades at that point in time all occurred in food chains with algae as the primary producer. Trophic cascades, Strong argued, may only occur in communities with fast-growing producers which lack defenses against herbivory.

Subsequent research has documented trophic cascades in terrestrial ecosystems, including:
# In the coastal prairie of Northern California, yellow bush lupines are fed upon by a particularly destructive herbivore, the root-boring caterpillar of the ghost moth. Entomopathogenic nematodes kill the caterpillars, and can increase the survival and seed production of lupines [Strong DR, Whipple AV, Child AL, Dennis B (1999) Model selection for a subterranean trophic cascade: Root-feeding caterpillars and entomopathogenic nematodes. Ecology 80:2750-2761] [Preisser EL (2003) Field evidence for a rapidly cascading underground food web. Ecology 84: 869-874] .
# In Costa Rican rain forest, a Clerid beetle specializes in eating ants. The ant "Pheidole bicornis" has a mutualistic association with "Piper" plants: the ant lives on the "Piper" and removes caterpillars and other insect herbivores. The Clerid beetle, by reducing the abundance of ants, increases the leaf area removed from "Piper" plants by insect herbivores [Letourneau DK, Dyer LA (1998) Experimental test in lowland tropical forest shows top-down effects through four trophic levels Ecology 79:1678-1687]

Critics pointed out, however, that published terrestrial trophic cascades generally involved smaller subsets of the food web (often only a single plant species). This was quite different from aquatic trophic cascades, in which the biomass of producers as a whole were reduced when predators were removed. Additionally, most terrestrial trophic cascades did not demonstrate reduced plant biomass when predators were removed, but only increased plant damage from herbivores [Polis GA, Sears ALW, Huxel GR, et al. (2000) When is a trophic cascade a trophic cascade? Trends in Ecology & Evolution 15: 473-475] . It was unclear if such damage would actually result in reduced plant biomass or abundance. In a recent meta-analysis, trophic cascades were generally "weaker" in terrestrial ecosystems, meaning that changes in predator biomass resulted in smaller changes in plant biomass [Shurin JB, Borer ET, Seabloom EW, Anderson K, Blanchette CA, Broitman B, Cooper SD, Halpern BS (2002) A cross-ecosystem comparison of the strength of trophic cascades. Ecological Letters 5:785-791] .

See also

* Food web
* Soil food web
* Lake ecology
* Stream ecology
* Mesopredator release hypothesis
* Population dynamics of fisheries

References

Bibliography