Combined Effects of Heavy Metal and Simulated Herbivory on Leaf Trichome Density in Sunflowers

Trichomes play a key role in both heavy metal tolerance and herbivory defense, and both stressors have been shown to induce increased trichome density. However, the combined effect of these stressors on trichome density in general, and specifically on metal-hyperaccumulating plants, has yet to be examined. The aim of this study was to test the effect of cadmium availability and herbivory on leaf trichome density and herbivore deterrence in the metal hyperaccumulator Helianthus annuus. To test this, H. Annuus plants were grown in control pots or pots inoculated with 10 mg/kg cadmium and were subjected to either no herbivory or simulated herbivory using mechanical damage and foliar jasmonic acid application. Herbivore deterrence was tested in a feeding assay using Spodoptera littoralis caterpillars. Interestingly, while the trichome density of H. annuus increased by 79% or 53.5% under high cadmium availability or simulated herbivory, respectively, it decreased by 26% when the stressors were combined. Furthermore, regardless of cadmium availability, simulated herbivory induced a 40% increase in deterrence of S. littoralis. These findings suggest that the combination of metal availability and herbivory might present excessive stress to hyperaccumulators. Moreover, they suggest that the risk of metal bioaccumulation in phytoremediation can be reduced by simulated herbivory.