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In ecology, the competitive exclusion principle, sometimes referred to as Gause's law, is a proposition that two species which compete for the same limited resource cannot coexist at constant population values. When one species has even the slightest advantage over another, the one with the advantage will dominate in the long term. This leads either to the extinction of the weaker competitor or to an evolutionary or behavioral shift toward a different ecological niche. The principle has been paraphrased in the maxim "complete competitors can not coexist". The competitive exclusion principle is classically attributed to Georgy Gause, although he actually never formulated it. The principle is already present in Darwin's theory of natural selection. Throughout its history, the status of the principle has oscillated between a priori ('two species coexisting must have different niches') and experimental truth ('we find that species coexisting do have different niches'). Based on field observations, Joseph Grinnell formulated the principle of competitive exclusion in 1904: "Two species of approximately the same food habits are not likely to remain long evenly balanced in numbers in the same region. One will crowd out the other". Georgy Gause formulated the law of competitive exclusion based on laboratory competition experiments using two species of Paramecium, P. aurelia and P. caudatum. The conditions were to add fresh water every day and input a constant flow of food. Although P. caudatum initially dominated, P. aurelia recovered and subsequently drove P. caudatum extinct via exploitative resource competition. However, Gause was able to let the P. caudatum survive by differing the environmental parameters (food, water). Thus, Gause's law is valid only if the ecological factors are constant. Gause also studied competition between two species of yeast, finding that Saccharomyces cerevisiae consistently outcompeted Schizosaccharomyces kefir by producing a higher concentration of ethyl alcohol.
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