Grazing tolerance of Gentianella amarella and other monocarpic herbs: why is tolerance highest at low damage levels?

Plants have adapted to compensate for the loss of vegetative biomass and reproductive potential caused by grazing. Shoot damage breaks down the correlative inhibition maintained by apical dominance. The consequent increased branching may lead to increased production of flowers and fruits in damaged plants, provided that enough resources, both in terms of meristems and nutrients, are available. In Gentianella amarella, the removal of the apex of the main stem (10% clipping) had no pronounced effect on branching and plant performance. In one of the two study populations, however, apically damaged plants produced more fruits than undamaged control plants. The plants also fully compensated for 50% removal of the main stem in terms of above-ground biomass, but their fruit production was reduced compared to control and apically damaged plants. After 75% clipping, fruit production was not significantly reduced compared to 50% clipping. Consequently, G. amarella showed highest tolerance in the presence of minor shoot damage. The pattern is qualitatively similar in some other monocarpic species (Gentianella campestris, Erysimum strictum and Rhinanthus minor). Multiple constraints as well as selective forces may shape these compensatory responses: (1) A lack of basal meristems may constrain tolerance of high damage levels. (2) Species with basal meristems may have a potential to tolerate major damage, but a shortage of resources or otherwise unfavourable growth conditions may constrain their compensatory ability. (3) It may be adaptive to have maximum tolerance of low and moderate damage levels if chemical defences reduce the risk of extensive shoot damage as well as the risk of repeated grazing. (4) The compensatory ability of monocarpic species may be affected by selective forces that favour fast vertical growth early in the season and unbranched architecture in undamaged conditions. Therefore, it is not the mere grazing history, but also other factors associated with growth conditions that are required to explain the variation in grazing tolerance.