Reproduction alternation in trees: testing the resource depletion hypothesis using experimental fruit removal in Quercus ilex

The keystones of resource budget models to explain mast seeding are that fruit production depletes tree stored resources, which become subsequently limiting to flower production the following year. These two hypotheses have, however, rarely been tested in forest trees. Using a fruit removal experiment, we tested whether preventing fruit development would increase nutrient and carbohydrates storage and modify allocation to reproduction and vegetative growth the following year. We removed all the fruits from nine adult Quercus ilex L. trees shortly after fruit set and compared, with nine control trees, the concentrations of nitrogen (N), phosphorus (P), zinc (Zn), potassium (K) and starch in leaves, twigs and trunk before, during and after the development of female flowers and fruits. The following year, we measured the production of vegetative and reproductive organs as well as their location on the new spring shoots. Fruit removal prevented the depletion of N and Zn in leaves during fruit growth. It also modified the seasonal dynamics in Zn, K and starch in twigs, but had no effect on reserves stored in the trunk. Fruit removal increased the production of female flowers and leaves the following year, and decreased the production of male flowers. Our results show that resource depletion operates differently for male and female flowering, because the timing of organ formation and the positioning of flowers in shoot architecture differ between male and female flowers. Our results suggest that N and Zn availability constrain flower production in Q. ilex, but also that other regulatory pathways might be involved. They strongly encourage further experiments manipulating fruit development over multiple years to describe the causal relationships between variations in resource storage and/or uptake, and male and female flower production in masting species.