- R/K selection theory
ecology, r/K selection theory relates to the selection of traits which promote success in particular environments. The theory originates from work on island biogeographyby the ecologists Robert MacArthurand E. O. Wilson[MacArthur, R. and Wilson, E. O. (1967). " The Theory of Island Biogeography", Princeton University Press (2001 reprint), ISBN 0-691-08836-5M.] .
In r/K selection theory, selective pressures are
hypothesised to drive evolutionin one of two generalized directions: "r"- or "K"-selection [Pianka, E. R. (1970). On r and K selection. "American Naturalist" 104, 592-597.] . These terms, r and K, are derived from standard ecological algebra, as illustrated in the simple of population dynamics[Verhulst, P. F. (1838). Notice sur la loi que la population pursuit dans son accroissement. "Corresp. Math. Phys." 10, 113-121.] :
where "r" is the growth rate of the
population("N"), and "K" is the carrying capacityof its local environmental setting. Typically, r-selected species exploit empty niches, and produce many offspring, each of which has a relatively low probability of surviving to adulthood. In contrast, K-selected species are strong competitors in crowded niches, and invest more heavily in fewer offspring, each of which has a relatively high probability of surviving to adulthood. In the scientific literature, r-selected species are occasionally referred to as "opportunistic", while K-selected species are described as "equilibrium" [For example: Weinbauer, M.G. and Höfle, M.G. (1998). [http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=106546 Distribution and Life Strategies of Two Bacterial Populations in a Eutrophic Lake] . "Appl. Environ. Microbiol." 64, 3776–3783.] .
r/K selection and environmental stability
In unstable or unpredictable environments r-selection predominates, as the ability to reproduce quickly is crucial, and there is little advantage in adaptations that permit successful competition with other organisms, because the environment is likely to change again. Traits that are thought to be characteristic of r-selection include: high
fecundity, small body size, early maturity onset, short generation time, and the ability to disperse offspring widely. Organisms whose life history is subject to r-selection are often referred to as r-strategists or r-selected. Organisms with r-selected traits range from bacteriaand diatoms, through insectsand weeds, to various semelparous cephalopods and mammals, especially small rodents.
In stable or predictable environments K-selection predominates, as the ability to compete successfully for limited resources is crucial, and populations of K-selected organisms typically are very constant and close to the maximum that the environment can bear. Traits that are thought to be characteristic of K-selection include: large body size, long
life expectancy, and the production of fewer offspring that require extensive parental care until they mature. Organisms whose life history is subject to K-selection are often referred to as K-strategists or K-selected. Organisms with K-selected traits include large organisms such as elephants, humansand whales, but smaller organisms also use this "strategy" successfully, such as Arctic Terns.
r/K selection as a continuous spectrum
It should be noted that, although some organisms are primarily r- or K-strategists, the majority of organisms fall between these two ecological extremes and may display traits considered characteristic of both ends of the r/K spectrum. For instance, trees have traits such as longevity and strong competitiveness that characterise them as K-strategists. In reproduction, however,
trees typically produce thousands of offspring and disperse them widely, traits characteristic of r-strategists. Similarly, reptiles such as sea turtles display both r- and K-traits: although large organisms with long lifespans (should they reach adulthood), they produce large numbers of unnurtured offspring.
r/K selection and ecological succession
In areas of major ecological disruption or sterilisation (such as after a major volcanic eruption, as at
Krakatoaor Mount Saint Helens), r- and K-strategists play distinct roles in the ecological successionthat regenerates the ecosystem. Because of their higher reproductive rates and ecological opportunism, primary colonisers typically are r-strategists and they are followed by a succession of increasingly competitive floraand fauna. The ability of an environment to increase energetic content, through photosynthetic capture of solar energy, increases with the increase in complex biodiversityas r species proliferate to reach a peak possible with K strategies [Gunderson, Lance & Holling, C. S. (Eds) (2001), "Panarchy: Understanding Transformations in Human and Natural Systems" (Island Press)] . Eventually a new equilibrium is approached (sometimes referred to as a climax community), with r-strategists gradually being replaced by K-strategists which are more competitive and better adapted to the emerging micro-environmental characteristics of the landscape. Typically, biodiversity is maximised at this stage, with introductions of new species resulting in the replacement and local extinctionof endemic species [McNeely, J. A. (1994) [http://www.springerlink.com/content/t76125571tr97m64/ Lessons of the past: Forests and Biodiversity] . "Biodiversity and Conservation" 3, 3-20.] .
tatus of r/K selection theory
Although r/K selection theory became widely used during the 1970s [Gadgil, M. and Solbrig, O.T. (1972). Concept of r-selection and K-selection - evidence from wild flowers and some theoretical consideration. "Am. Nat." 106, 14.] [Long, T. and Long, G. (1974). Effects of r-selection and K-selection on components of variance for 2 quantitative traits. "Genetics" 76, 567-573.] [Grahame, J. (1977). Reproductive effort and r-selection and K-selection in 2 species of "Lacuna" (Gastropoda-Prosobranchia). "Mar. Biol." 40, 217-224.] [Luckinbill, L.S. (1978). r and K selection in experimental populations of "Escherichia coli". "Science" 202, 1201-1203.] , it also began to attract more critical attention [Barbault, R. (1987). Are still r-selection and K-selection operative concepts? "Acta Oecologica-Oecologia Generalis" 8, 63-70.] [Kuno, E. (1991). Some strange properties of the logistic equation defined with r and K - inherent defects or artifacts. "Researches on Population Ecology" 33, 33-39.] [Getz, W.M. (1993). Metaphysiological and evolutionary dynamics of populations exploiting constant and interactive resources - r-K selection revisited. "Evolutionary Ecology" 7, 287-305.] . In particular, an influential review by the ecologist
Stephen Stearnsdrew attention to gaps in the theory, and to ambiguities in the interpretation of empirical data for testing it [cite journal|author=Stearns, SC|year=1977|title=Evolution of life-history traits - critique of theory and a review of data|journal=Ann. Rev. of Ecology and Systematics|volume=8|pages=145-171|url=http://faculty.washington.edu/kerrb/Stearns1977.pdf|format=PDF] .
Evolutionary game theory
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