Plant architecture, sectoriality and plant tolerance to herbivores

The evolution of tolerance is one potential plant response to selection imposed by herbivores. Plant architecture, and in turn, sectoriality may influence a plant's ability to tolerate tissue loss. However, each may either constrain or facilitate a plant's ability to compensate following herbivore attack depending on the plant part damaged and the identity of the damaging herbivore. Plants are limited in their ability to respond to localized damage by chewing insects because carbon does not flow freely from damaged to undamaged plant parts, particularly between branches. Thus, defoliation of individual branches invariably results in decreased growth and reproduction of those branches. Within branches, carbon flow via vascular connections between orthostichies may ameliorate the effects of damage restricted within an orthostichy. Local induction of secondary chemicals to spread damage by folivores throughout a plant's canopy, redistribution of resources within and between IPU's, and delaying reproductive activity until resources have been pooled may all alleviate the constraints on response of plants to grazing. In contrast to the effects of damage by grazers, the metameric construction of plants typically ensures points of regrowth from dormant buds when apical meristems are destroyed either by vertebrate browsers or galling insects. Sectoriality constrains the ability of sap-sucking insects to tap the entire resource base of a plant, thus having a positive effect on plant fitness. However, both the site and timing of attack mitigate the degree of limitation imposed by sectoriality. During peak periods of assimilation, photosynthate flow is mainly over short distances (between sources and sinks within the canopy), and thus sap-sucking insects have a small resource base to draw upon. In contrast, when sucking insects tap into vascular elements in which the flow is from roots to leaves and vice versa, resource availability to the insect (and in turn, potential resource loss from the plant) are only limited by the resources present in those vascular elements. Studies of specific traits in species which demonstrate differential tolerance would greatly add to our understanding of herbivore impacts on plant growth and reproduction. In particular, intraspecific variation in tolerance has been documented for individuals within and among populations with different grazing histories. A number of traits related to sectoriality and architecture probably contribute to such variation in tolerance, and because they are easily manipulated and easily quantified, represent potentially profitable avenues of research. These traits include distribution of leaves and buds, ability to release secondary meristems from dormancy, and the timing of resource movement both before and subsequent to damage.