How to quantify plant tolerance to loss of biomass?

In some plant species the whole shoot is occasionally removed, as a result of specialist herbivory, grazing, mowing, or other causes. The plant can adapt to defoliation by allocating more to tolerance and less to growth and defense. Plant tolerance to defoliation (TOL1) is typically measured as the ratio between the average dry weight of a group of damaged plants and a control group of undamaged plants, both measured some time after recovery. We develop a model to clarify what TOL1 actually measures. We advocate keeping regrowth (REG2) and shoot-root ratio, both elements of TOL1, separate in the analysis. Based on a resource trade-off, exotic plants from populations in the USA (no specialist herbivory) are expected to grow faster and be less tolerant than native Dutch populations (with specialist herbivory). Indeed Dutch plants had both a significantly larger fraction biomass in roots and faster regrowth (REG2), while US plants attained the highest weight in the control without defoliation. Using key-factor analysis, we illustrate how growth rates, regrowth, and shoot-root ratio each contribute to final biomass (plant fitness). Our proposed method gives more insight in the mechanisms that underly plant tolerance against defoliation and how tolerance contributes to fitness.