Nitrogen fertilizer and gender effects on the secondary metabolism of yaupon, a caffeine-containing North American holly
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Yaupon (Ilex vomitoria) is a caffeine-containing dioecious shrub native to the southeastern United States that was historically brewed into a stimulating beverage. We tested predictions of the carbon/nutrient balance (CNB) hypothesis by determining whether nitrogen availability and gender influence production of caffeine and related alkaloids as well as phenolic compounds in leaves of pot-grown yaupon plants fertilized with ammonium nitrate. The CNB hypothesis predicts that additional nitrogen should result in increased alkaloid concentrations and decreased phenolic concentrations. An extension of the CNB hypothesis to dioecious plants predicts that females have higher C/N ratios and therefore higher phenolic concentrations and lower alkaloid concentrations than male conspecifics. In our study, caffeine and total alkaloid concentrations were 5–10 times higher in fertilized than control plants but did not vary by gender. Nevertheless, an observed interaction between gender and fertilization suggests that females respond more to fertilization than males in caffeine production. In addition, fertilized plants not only contained higher concentrations of alkaloids and total nitrogen but also allocated a larger proportion of their nitrogen to alkaloid production than control plants. Total phenolic concentrations were higher in control females than control males as predicted by the CNB hypothesis, but did not vary by treatment nor were there differences by gender among fertilized plants. We also found high correlations between antioxidant capacity and both classes of phenolic compounds detected in our study (cinnamic acid derivatives and flavonoids) indicating that in addition to their putative defensive function against herbivores, phenolics protect yaupon from oxidative stress. Explanation of the inconsistencies between our data and predictions of the CNB hypothesis may benefit from a re-appraisal of the physiological mechanisms by which resource availability affects secondary metabolism as well as consideration of the selective pressures to which secondary metabolism responds.