Evolution of Plants : a mathematical perspective

Sammanfattning: The Earth harbors around 300 000 plant species. The rich and complex environment provided by plants is considered a key factor for the extraordinary diversity of the terrestrial fauna by, for example, providing food and shelter. This thesis contributes to the understanding of these questions by investigating how the interplay of physiology, demography, and evolution gives rise to variation and diversity in fundamental plant traits. This will help us answer questions such as: How has this amazing diversity of plant species emerged? Which mechanisms maintain diversity? How are plant strategies and plant diversity influenced by variations in the environment?A plant faces multiple problems to survive and reproduce successfully. These problems can be modeled by considering traits, trade-offs and a fitness measure. For example: How to maximize growth rate, while maximizing structural stability? I will investigate four plant models in order to understand the function of plants, and mechanisms promoting diversity. Paper I: We study how annual plants with and without growth constraints should optimize their flowering time when productivity or season length changes. With a dynamic ontogenetic growth model and optimal control theory we prove that a bang-bang reproductive control is optimal under constrained growth and constant mortality rate. We find that growth constraints can flip the direction of optimal phenological response for increasing productivity. The reason is that the growth rate of vegetative mass saturates at high productivity and therefore it is better to flower earlier and take advantage of a longer reproductive period. If season length extends equally both in the beginning and the end of the season, growth constraints control the direction of the optimal response as well. Our theory can help explaining phenological patterns along productivity gradients, and can be linked to empirical observations made on a calendar scale.Paper II: We introduce a new measure of tree crown-rise efficiency based on the loss of biomass of the tree during growth. The more mass the tree looses during growth, the less crown-rise efficient it is. Top-heavy shapes loose more biomass than bottom-heavy shapes. Light-use efficiency is defined as the mean light assimilation of the leaves in the crown times the ratio of leaf mass and total mass. We then study the trade-off between light-use efficiency to crown-rise efficiency for tree crown shapes. We assume that the total tree mass is constant, and a constant vertical light gradient represent the shading from a surrounding forest. We find large differences in crown shapes at intermediate vertical light gradient, when both self-shading and mean-field shading are important, suggesting light-use vs crown-rise efficiency as a new trade-off that can explain tree diversity. Our crown-rise efficiency measure could easily be integrated into existing forest models.Paper III: We extend an evolutionary tree crown model, where trees with different heights compete for light, with drought-induced mortality rates depending on ground-water availability and the depth of an optional taproot. The model does not include competition for ground water. Our model explains how ground-water availability can shape plant communities, when taproot and non-taproot strategies can coexist, and when only one of these strategies can persist. We investigate how emerging plant diversity varies with water table depth, soil water gradient and drought-induced mortality rate. The taproot enables plants to reach deep water, thus reducing mortality, but also carries a construction cost, thus inducing a trade-off. We find that taproots maintain plant diversity under increasing drought mortality, and that taproots evolve when groundwater is accessible at low depths. There are no viable strategies at high drought mortality and deep water table. Red Queen evolutionary dynamics appear at intermediate drought mortality in mixed communities with and without taproots, as the community never reaches a final evolutionarily stable composition.Paper IV: We extend a size-structured plant model, with self-shading and two evolving traits, crown top-heaviness and crown width-to-height ratio. The model allows us to identify salient trade-offs for the crown shape. The most important trade-off for top-heaviness is light-use efficiency vs crownrise efficiency, and the most important trade-off for width-to-height ratio is self-shading vs branch costs. We find that when the two traits coevolve; the outcome is a single common evolutionarily stable strategy (ESS), far away from the highest net primary production (NPP). When only sun angle is decreasing with increasing latitude both the crown width-to-height ratio and crown top-heaviness decrease. However, when light response in addition to the sun angle decreases with increasing latitude, the crown width-to-height ratio is nearly invariant of latitude except at low site productivity when the ratio decreases with latitude. Top-heaviness is always decreasing with increasing latitude. Finally, we find that crown top-heaviness increases with the NPP or leaf-area index (LAI) at ESS, but crown width-to-height ratio is maximal at an intermediate NPP or LAI.