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Assuming a stationary distribution of environmental states with no temporal autocorrelation, a diffusion approximation for population size and allele frequency,<jats:italic>p<\/jats:italic>, reveals that the expected change in<jats:italic>p<\/jats:italic>involves the gradient with respect to<jats:italic>p<\/jats:italic>of the stochastic intrinsic rate of increase (the density\u2010independent long\u2010run growth rate),<jats:inline-graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" xlink:href=\"graphic\/JEB_1533_mu1.gif\" xlink:title=\"inline image\"\/>, where<jats:italic>r<\/jats:italic>is the mean Malthusian fitness in the average environment and<jats:inline-graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" xlink:href=\"graphic\/JEB_1533_mu2.gif\" xlink:title=\"inline image\"\/>is the environmental variance in population growth rate. The expected relative fitness of a genotype is its Malthusian fitness in the average environment minus the covariance of its fitness with population growth rate. The influence of fitness correlation between genotypes is illustrated by an analysis of the Haldane\u2013Jayakar model of fluctuating selection on a single diallelic locus, and on two loci with additive effects on a quantitative character.<\/jats:p>","DOI":"10.1111\/j.1420-9101.2008.01533.x","type":"journal-article","created":{"date-parts":[[2008,4,15]],"date-time":"2008-04-15T01:50:24Z","timestamp":1208224224000},"page":"1096-1105","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":31,"title":["Adaptive topography of fluctuating selection in a Mendelian population"],"prefix":"10.1093","volume":"21","author":[{"given":"R.","family":"LANDE","sequence":"first","affiliation":[]}],"member":"286","published-online":{"date-parts":[[2008,4,14]]},"reference":[{"key":"e_1_2_6_2_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-642-93128-4"},{"key":"e_1_2_6_3_1","doi-asserted-by":"publisher","DOI":"10.1023\/A:1013373907708"},{"key":"e_1_2_6_4_1","doi-asserted-by":"publisher","DOI":"10.1017\/S0016672300023156"},{"key":"e_1_2_6_5_1","volume-title":"The Mathematical Theory of Selection, Recombination, and Mutation","author":"B\u00fcrger R.","year":"2000"},{"key":"e_1_2_6_6_1","doi-asserted-by":"publisher","DOI":"10.1093\/genetics\/152.2.807"},{"key":"e_1_2_6_7_1","volume-title":"Matrix Population Models","author":"Caswell H.","year":"2001"},{"key":"e_1_2_6_8_1","doi-asserted-by":"publisher","DOI":"10.1017\/CBO9780511525711"},{"key":"e_1_2_6_9_1","doi-asserted-by":"crossref","first-page":"462","DOI":"10.2307\/1426109","article-title":"Ergodicity of age structure in populations with Markovian vital rates. 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