D. Pilson, The evolution of plant response to herbivory: simultaneously considering resistance and tolerance in Brassica rapa, EVOL ECOL, 14(4-6), 2000, pp. 457-489
Although the evolution of plant response to herbivory can involve either re
sistance (a decrease in susceptibility to herbivore damage) or tolerance (a
decrease in the per unit effect of herbivory on plant fitness), until rece
ntly few studies have explicitly incorporated both of these characters. Mor
eover, theory suggests these characters do not evolve independently, and al
so that the pattern of natural selection acting on resistance and tolerance
depends on their costs and benefits. In a genotypic selection analysis on
an experimental population of Brassica rapa (Brassicaceae) I found a comple
x set of correlational selection gradients acting on resistance and toleran
ce of damage by flea beetles (Phyllotreta cruciferae: Chrysomelidae) and we
evils (Ceutorhynchus assimilis: Curculionidae), as well as directional and
stabilizing selection on resistance to attack by weevils. Evolution of resp
onse to flea beetle attack is constrained by a strong allocation cost of to
lerance, and this allocation cost may be caused by a complex correlation am
ong weevil resistance, weevil tolerance, flea beetle resistance, and flea b
eetle tolerance. Thus, one important conclusion of this study is that ecolo
gical costs may involve complex correlations among multiple characters, and
for this reason these costs may not be detectable by simple pairwise corre
lations between characters. The evolution of response to weevil attack is p
robably constrained by a series of correlations between weevil resistance,
weevil tolerance, and fitness in the absence of weevil damage, and possibly
by a cost of tolerance of weevil damage. However, the nature of these cons
traints is complicated by apparent overcompensation for weevil damage. Beca
use damage by both flea beetles and weevils had non-linear effects on plant
fitness, standard measures of tolerance were not appropriate. Thus, a seco
nd important contribution of this study is the use of the area under the cu
rve defined by the regression of fitness on damage and damage-squared as a
measure of tolerance.