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Trophic cascades occur when predators in a food web suppress the abundance or alter the behavior of their prey, thereby releasing the next lower trophic level from predation (or herbivory if the intermediate trophic level is a herbivore). For example, if the abundance of large piscivorous fish is increased in a lake, the abundance of their prey, smaller fish that eat zooplankton, should decrease. The resulting increase in zooplankton should, in turn, cause the biomass of its prey, phytoplankton, to 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.
A Top Down Cascade is a trophic cascade where the food chain or food web is disrupted by the removal of a top predator, or a third or fourth level consumer. On the other hand, a bottom up cascade occurs when a primary producer, or primary consumer is removed, and there is a reduction of population size through the community. An example could include Paine's study from the University of Washington, where he removed several species in different plots along the North-Western United States coast line, and realized that Pisaster, a common starfish, when removed, created a top-down cascade which involved a surge in barnacle and mussel populations, but a decrease in the populations of chitons, limpets, and whelks. This led to the conclusion that Pisaster was a keystone species in that food web.
Aldo Leopold is generally credited with first describing the mechanism of a trophic cascade, based on his observations of overgrazing of mountain slopes by deer after human extermination of wolves. Nelson Hairston, Frederick E. Smith and Lawrence B. Slobodkin are generally credited with introducing the concept into scientific discourse, although they did not use the term either. Hairston, Smith and Slobodkin argued that predators reduce the abundance of herbivores, allowing plants to flourish. 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 (e.g. 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 (e.g. 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.
Hairston, Smith and Slobodkin argued that the ecological communities acted as food chains with three trophic levels. Subsequent models expanded the argument to food chains with more than or fewer than three trophic levels. 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.
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:
Antagonistically, this principle is sometimes called the "trophic trickle".
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:
A classic example of a terrestrial tropic cascade is the reintroduction of gray wolves (Canis lupus) to Yellowstone National Park, which reduced the number and behavior of elk (Cervus elaphus). This in turn released several plant species from grazing pressure and subsequently led to the transformation of riparian ecosystems. This example of a trophic cascade is vividly shown and explained in a viral video “How Wolves Change Rivers.”
The fact that the earliest documented trophic cascades all occurred in lakes and streams led Donald Strong to speculate that fundamental differences between aquatic and terrestrial food webs made trophic cascades primarily an aquatic phenomenon. 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:
Critics pointed out 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. It was unclear if such damage would actually result in reduced plant biomass or abundance. In 2002 a meta-analysis found trophic cascades to be generally weaker in terrestrial ecosystems, meaning that changes in predator biomass resulted in smaller changes in plant biomass. In contrast, a study published in 2009 demonstrated that multiple species of trees with highly varying autecologies are in fact heavily impacted by the loss of an apex predator. Another study, published in 2011, demonstrated that the loss of large terrestrial predators also significantly degrades the integrity of river and stream systems, impacting their morphology, hydrology, and associated biological communities.
The critics' model is challenged by studies accumulating since the reintroduction of gray wolves (Canis lupus) to Yellowstone National Park. The gray wolf, after being extirpated in the 1920s and absent for 70 years, was reintroduced to the Park in 1995 and 1996. Since then a three-tiered trophic cascade has been reestablished involving wolves, elk (Cervus elaphus), and woody browse species such as aspen (Populus tremuloides), cottonwoods (Populus spp.), and willows (Salix spp.). Mechanisms likely include actual wolf predation of elk, which reduces their numbers, and the threat of predation, which alters elk behavior and feeding habits, resulting in these plant species being released from intensive browsing pressure. Subsequently, their survival and recruitment rates have significantly increased in some places within Yellowstone's northern range. This effect is particularly noted among the range's riparian plant communities, with upland communities only recently beginning to show similar signs of recovery.
Examples of this phenomenon include:
Trophic cascades also impact the biodiversity of ecosystems, and when examined from that perspective wolves appear to be having multiple, positive cascading impacts on the biodiversity of Yellowstone National Park. These impacts include:
There are a number of other examples of trophic cascades involving large terrestrial mammals, including:
In addition to the classic examples listed above, more recent examples of trophic cascades in aquatic systems have been identified:
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