Genetic and environmental variation of functional traits within populations might be maintained by natural selection when resource allocation costs (RACs) balance trait benefits. Despite the intuitive appeal of optimization models, empirical tests have failed to support the importance of RACs for plant traits that confer resistance against pests. To address this discrepancy, we modified an early model by allowing the cost function to vary across a resource gradient as predicted for RACs and by assuming that the benefits depend on variation in the pest population for susceptibility. Instead of the intermediate defense optimum of the original model, defenses were predicted to be either high or absent, depending on resource availability and history. This result is not supported by empirical tests for ecological or evolutionary outcomes, including our own examination of glucosinolate toxins from sib-families of Boechera stricta (Brassicaceae) grown across an NPK fertilizer gradient. Although we detected an apparent cost of defense in the absence of herbivores, the cost did not increase as resources became more limiting. Also defense production did not vary across the resource gradient as predicted by the modified model. Thus, a model based on explicit expectations of RACs produced predictions that are not supported. Instead, other kinds of costs, such as ecological (indirect) costs may be more important. Alternatively, general conflicts in gene expression and antagonistic crosstalk among signaling pathways may underlie apparent costs.

Reports of forest damage have increased with the frequency of climatic extremes, but longer term impacts of such events on population dynamics of forest trees are generally unknown. Incited by the turn-of-the-century drought, sudden aspen decline (SAD) damaged 535 000 ha of Populus tremuloides Michx. in the Southern Rockies ecoregion of western North America. Although spread of the disease stopped in about 2009, most of the affected stands continued to deteriorate. Remeasurement of plots in southwestern Colorado showed that, since the peak of the epidemic, live basal area in sick plots decreased by an additional 28% to only 38% of that in healthy plots. Sick plots had much more recent damage than healthy plots, with almost three times as much recently dead basal area, over twice the density of recently dead trees, and almost four times as much recent crown loss. The important contributing agents in SAD were still active in sick stands in 2013. Density of small regeneration showed opposite trends, increasing in healthy plots and decreasing in sick plots. Timely regeneration treatments may be needed in some such stands to facilitate recovery. In addition to acute damage from climatic extremes, long-term decline diseases like SAD will likely be a common signature of forest damage from climate change.