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Mutualism

[myoo-choo-uh-liz-uhm]

Mutualism is a biological interaction between individuals of two different species, where both individuals derive a fitness benefit, for example increased survivorship. Similar interactions within a species are known as co-operation. It can be contrasted with interspecific competition, in which both species experience reduced fitness, and exploitation, in which one species benefits at the expense of the other. Mutualism and symbiosis are sometimes used as if they are synonymous, but this is strictly incorrect: symbiosis is a broad category, defined to include relationships which are mutualistic, parasitic or commensal. Mutualism is therefore only one type. Mutualism plays a key part in ecology and evolutionary biology. For example, mutualistic interactions are vital for terrestrial ecosystem function as more than 70% of land plants rely on mycorrhizal relationships with fungi to provide them with inorganic compounds and trace elements. In addition, mutualism has driven the evolution of much of the biological diversity we see, such as flower forms (important for pollination mutualisms) and co-evolution between groups of species. However mutualism has historically received less attention than other interactions such as predation and parasitism.

Measuring the exact fitness benefit to the individuals is not always straightforward, particularly when the individuals can receive benefits from a range of species, for example most plant-pollinator mutualisms. It is therefore common to categorise mutualisms according to the closeness of the association, using terms such as obligate versus facultative. Defining "closeness", however, is also problematical. It can refer to mutual dependency (the species cannot live without one another) or the biological intimacy of the relationship in relation to physical closeness (e.g. one species living within the tissues of the other species).

Types of relationships

Mutualistic interactions can be thought of as a form of "biological barter in which species trade resources (for example carbohydrates or inorganic compounds) or services such as gamete, offspring dispersal, or protection from predators.

Resource-resource relationships

Resource-resource interactions, in which one type of resource is traded for a different resource, are probably the most common form of mutualism; for example mycorrhizal associations between plant roots and fungi, with the plant providing carbohydrates to the fungus in return for primarily phosphate but also nitrogenous compounds. Other examples include rhizobia bacteria which fix nitrogen for leguminous plants (family Fabaceae) in return for energy-containing carbohydrates.

Service-resource relationships

Service-resource relationships are also common, for example pollination in which nectar or pollen (food resources) are traded for pollen dispersal (a service) or ant protection of aphids, where the aphids trade sugar-rich honeydew (a by-product of their mode of feeding on plant sap) in return for defence against predators such as ladybird beetles.

Service-service relationships

Strict service-service interactions are very rare, for reasons that are far from clear. One example is the relationship between sea anemones and anemonefish in the family Pomacentridae: the anemones provide the fish with protection from predators (which cannot tolerate the stings of the anemone's tentacles) and the fish defend the anemones against butterfly fish (family Chaetodontidae) which eat anemones. However, in common with many mutualisms, there is more than one aspect to the biological barter: in the anemonefish-anemone mutualism, waste ammonia from the fish feed the symbiotic algae that are found in the anemone's tentacles. Therefore what appears to be a service-service mutualism in fact has a service-resource component. A second example is that of the relationship between some ants and trees in the genus Acacia, such as the Whistling Thorn and Bullhorn Acacia. The ants nest inside the plant's thorns. In exchange for shelter, the ants protect acacias from attack by herbivores (which they frequently eat, introducing a resource component to this service-service relationship) and competition from other plants by trimming back vegetation that would shade the acacia. In addition, another service-resource component is present, as the ants regularly feed on lipid-rich food-bodies called Beltian bodies that are on the Acacia plant.

Humans and mutualism

Humans also engage in mutualisms with other species, including our gut flora (without which we would not be able to digest food efficiently) and domesticated animals such as horses, which provide transportation in return for food and shelter. In traditional agriculture, many plants will function mutualistically as companion plants, providing each other with shelter, soil fertility and the repelling of pests. For example, beans may grow up cornstalks as a trellis, while fixing nitrogen in the soil for the corn, as exploited in the Three Sisters gardening technique. The question how and why species might cooperate has been addressed philosophically by a number of writers. Gilles Deleuze, for example, was interested in the way this questioned the conception of evolutionism and the notion of linear historical progress.

References

Specific

General

Breton, Lorraine M., and John F. Addicott. 1992. Density-Dependent Mutualism in an Aphid-Ant Interaction. Ecology, Vol. 73, No. 6, pp. 2175-2180.
Bronstein, JL. 1994. Our current understand of mutualism. Quarterly Review of Biology 69 (1): 31-51 MAR 1994
Bronstein JL, 2001. The exploitation of mutualisms. Ecology Letters 4 (3): 277-287
Bronstein JL, 2001. The costs of mutualism. American Zoologist 41 (4): 825-839 S
Bronstein JL, Alarcon R, Geber M. 2006. The evolution of plant-insect mutualisms. New Phytologist 172 (3): 412-428
Denison RF, Kiers ET 2004. Why are most rhizobia beneficial to their plant hosts, rather than parasitic? Microbes and Infection 6 (13): 1235-1239
DeVries, PJ; and Baker, I. 1989. Butterfly exploitation of an ant-plant mutualism: Adding insult of herbivory. Journal of the New York Entomological Society [J. N.Y. ENTOMOL. SOC.]. Vol. 97, no. 3, pp. 332-340.
Hoeksema, J.D. & E.M.Bruna. 2000. Pursuing the big questions about interspecific mutualism: a review of theoretical approaches. Oecologia 125:321-330
Jahn, G.C. and J.W. Beardsley 2000. Interactions of ants (Hymenoptera: Formicidae) and mealybugs (Homoptera: Pseudococcidae) on pineapple. Proceedings of the Hawaiian Entomological Society 34: 181-185.
Jahn, Gary C., J. W. Beardsley and H. González-Hernández 2003. A review of the association of ants with mealybug wilt disease of pineapple. Proceedings of the Hawaiian Entomological Society. 36:9-28.
Noe, R. & P. Hammerstein. 1994. Biological markets: supply and demand determine the effect of partner choice in cooperation, mutualism and mating. Behavioral Ecology and Sociobiology 35:1-11
Ollerton, J. 2006. "Biological Barter": Patterns of Specialization Compared across Different Mutualisms. pp. 411-435 in: Waser, N.M. & Ollerton, J. (Eds) Plant-Pollinator Interactions: From Specialization to Generalization. University of Chicago Press.
Paszkowski, U. 2006. Mutualism and parasitism: the yin and yang of plant symbioses. Current Opinion on Plant Biology 9 (4): 364-370.
Porat, D. & Chadwick-Furman, N. E. 2004 Effects of anemonefish on giant sea anemones:expansion behavior,growth, and survival. Hydrobiologia 530, 513–520. (doi:10.1007/s10750-004-2688-y)
Porat, D. & Chadwick-Furman, N. E. 2005 Effects of anemonefish on giant sea anemones: ammonium uptake,zooxanthella content and tissue regeneration. Mar. Freshw. Behav. Phys. 38, 43–51. (doi:10.1080/102362405000 57929)
Thompson, J. N. 2005 The geographic mosaic of coevolution. University of Chicago Press.