Seed Dispersal as a Maternally Influenced Character: Mechanistic Basis of Maternal Effects and Selection on Maternal Characters in an Annual Plant

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Abstract
Maternal influences on progeny characters affect phenotypic correlations between characters expressed in maternal and progeny generations and consequently influence evolutionary responses to selection. Net selection on maternally influenced characters depends on selection both on the progeny character and on the maternal characters that influence it. I used seed dispersal in Cakile edentula as a system in which to identify the mechanisms of environmentally mediated maternal effects and to determine how selection on maternal characters alters the adaptive value of dispersal. In C. edentula, maternal morphology responds to conspecific density experienced by the mother. Maternal morphology in turn affects offspring (seed) dispersal and density and thereby offspring morphology and fitness. I estimated the magnitude of density-mediated maternal effects on dispersal and identified their mechanism by characterizing the plasticity of maternal morphology to density. I also measured density-dependent selection on maternal characters that influence dispersal. Maternal plasticity to density was caused by both allometric and nonallometric variation in morphology, and this plasticity resulted in a negative correlation between maternal and progeny density. Such negative maternal effects are expected to retard responses to selection. Maternal morphology influenced maternal fitness, in part through the relationship of fitness to maternal plant size and in part through size-independent fitness effects. Maternal phenotypes that promote dispersal, and thereby increase progeny fitness, were associated with decreased maternal fitness. Selection on dispersal at the level of progeny favors increased dispersal; maternal influences on dispersal, however, not only cause a greatly reduced adaptive value of dispersal but lead to the prediction of a slower response to selection.