Effects of variation in carbon, nitrogen, and phosphorus molarity and stoichiometry on sex determination in the fern Ceratopteris richardii

Abstract

Carbon (C), nitrogen (N), and phosphorous (P) are needed by all organisms for basic biological processes. When an individual macronutrient is not accessible, nutrient limitation occurs. The stochiometric balance between multiple nutrients and individual concentrations are both vital for normal growth and development. Labile sex expression in plants is a phenotypic trait predicted to be sensitive to local nutrient conditions because males and females differ in their nutritional demands. We applied concepts from ecological stoichiometry to assess the effects of variation in individual nutrient concentration and multiple macronutrient stoichiometry on sexual development in the fern Ceratopteris richardii Brongn. Manipulation of N, P, and organic and inorganic C was expected to yield variation in the ratio of males to females, consistent with environmental sex-determination theory. Our results suggest nutrient stoichiometry, not strictly concentration, influences sex determination at ambient CO2. However, an early response to population density preempted nutrient effects in elevated CO2 environments with exogenous glucose, in which C. richardii gametophytes presumably grow naturally. Although sex determination is not nutrient-dependent in the latter environment, C:N in the dry mass of meristic gametophytes is influenced by the external nutrient context, suggesting sex determination takes place before abiotic environmental factors subsequently influence plant nutrient uptake.