Green World Hypothesis
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The green world hypothesis proposes that
predators Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
are the primary regulators of ecosystems: they are the reason the world is 'green', by regulating the
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
s that would otherwise consume all the greenery. It is also known as the HSS hypothesis, after Hairston, Smith and Slobodkin, the authors of the seminal paper on the subject. Although plenty of herbivores exist that would potentially diminish the vegetation of the world, many researchers find themselves asking the question of how biomass and biodiversity are able to be maintained. The natural order to allow for the persistence of all species and ecosystems requires an opposite force acting upon these herbivores. A system of checks and balances is proposed in allowing the flourishing of flora in various ecosystems, as suggested by the green world hypothesis. In addition to plant defense mechanisms, predators assist in the regulation of these herbivore population numbers, limiting the amount of vegetation that is consumed. Several ecosystems are characterized by a
trophic cascade Trophic cascades are powerful indirect interactions that can control entire ecosystems, occurring when a trophic level in a food web is suppressed. For example, a top-down cascade will occur if predators are effective enough in predation to redu ...
system, in which all levels interact and impact the persistence of one another (Siliman and Angelini, 2012). For example, the herbivores reduce plant populations, but are kept in check by carnivorous consumers that limit population growth beyond what's allotted given resource availability. The study of trophic cascades is highly important to understanding the green world hypothesis. One way that trophic cascades can impact ecosystems is through the limitation of net primary productivity, which determines energy flow, through various resources. This bottom-up approach results in the abundance of unpalatable plant species due to various environmental conditions (Siliman and Angelini, 2012). Additionally, energy in a given system can be determined by predators at the highest trophic level, or the carnivores that consume other carnivores. This top-down approach is characterized by high consumer densities, and in many cases, weedy plant systems, without strong initial defense mechanisms in place (Siliman and Angelini, 2012). These processes of the maintenance of trophic cascades often operate simultaneously. A general consensus is that trophic cascades tend to have a larger effect in aquatic ecosystems compared to terrestrial. However, overregulation in any of these communities has the potential to result in the degradation of the trophic cascade within the system, preventing growth across many species of all levels (2012).


History

The green world hypothesis was likely first proposed in a 1957 course by Frederick Edward Smith at the
University of Michigan The University of Michigan (U-M, U of M, or Michigan) is a public university, public research university in Ann Arbor, Michigan, United States. Founded in 1817, it is the oldest institution of higher education in the state. The University of Mi ...
. In 1960,
Nelson Hairston Nelson Hairston Sr. (16 October 1917 – 31 July 2008) was an American ecologist. Hairston is well known for his work in ecology and human disease. In the field of ecology he is famous for championing the idea of the trophic cascade, on which he pu ...
, Smith, and
Lawrence Slobodkin Lawrence Basil Slobodkin (June 22, 1928 – September 12, 2009) was an American ecologist and Professor Emeritus at the Department of Ecology and Evolution, Stony Brook University, State University of New York. He was one of the leading pioneers ...
published a paper laying out the green world hypothesis. The name HSS derives from the first letters of each of their surnames. Robert T. Paine did experiments in 1966 with ''
Pisaster ochraceus ''Pisaster ochraceus'', generally known as the purple sea star, ochre sea star, or ochre starfish, is a common seastar found among the waters of the Pacific Ocean. Identified as a keystone species, ''P. ochraceus'' is considered an important ind ...
'' which illustrated their role as a
keystone species A keystone species is a species that has a disproportionately large effect on its natural environment relative to its abundance. The concept was introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in main ...
in regulating ''
Mytilus californianus The California mussel (''Mytilus californianus'') is a large edible mussel, a marine bivalve mollusk in the family Mytilidae. This species is native to the west coast of North America, occurring from northern Mexico to the Aleutian Islands of ...
''. In this study, Paine observed that biodiversity and ecosystem persistence in intertidal zones were highly dependent upon sea star presence (Ceci, 2020). Although this is recognized as a keystone species, Paine’s research allowed for this discovery to be made, noticing that mussels, anemones, alga, among other species reduced quantities following the decrease in sea star numbers (Ceci, 2020). This is a strong example of the importance of the maintenance of the trophic cascade and suggests that top-down control is the primary regulatory factor in this system. James Estes and John Palmisano did similar experiments with otters, sea urchins, and kelp, where otter presence increased kelp presence in a
trophic cascade Trophic cascades are powerful indirect interactions that can control entire ecosystems, occurring when a trophic level in a food web is suppressed. For example, a top-down cascade will occur if predators are effective enough in predation to redu ...
. As discussed in this study, sea otters, natural consumers of sea urchins, have allowed for the flourishing of kelp forests. Without the assistance and presence of these otterns, sea urchin biomass would be allowed to increase, which would likely pose a significant threat to the ecosystem. The absence of sea otters in the ecosystem results in the near elimination of kelp forests, forming an ecological concept known as urchin barrens (Siliman and Angelini, 2012). John Terborgh examined Venezuelan valleys with and without predators in 2006, demonstrating the green world hypothesis on land.


Criticism

The plant self-defense hypothesis proposes that plants are not entirely consumed by herbivores primarily because of their adaptations against it (thorns, toxicity, cellulose, etc.).


References

{{Modelling ecosystems Trophic ecology