Disentangling dimensions of phytochemical diversity: alpha and beta have contrasting effects on an insect herbivore.

Phytochemical diversity is comprised of two main dimensions-the average (alpha) within-plant neighbors or the difference (beta) in the composition of chemicals between plant neighbors. Research, however, has primarily examined the consequences of phytochemical diversity on herbivore performance through a single dimension, even though diversity is multidimensional. Furthermore, the ecological role of phytochemical diversity is not well understood because each of these dimensions exhibits unique biological effects on herbivore performance. Therefore, it has been difficult to tease apart the relative importance of alpha and beta chemical diversities on plant-herbivore interactions. We experimentally manipulated alpha and beta diversities along a chemical gradient to disentangle the relative effects of these dimensions on the performance of a mobile generalist herbivore, Trichoplusia ni (Hübner), using 16 genotypes from the Solanum pennellii introgression lines. First, we found contrasting effects of alpha and beta diversities on herbivore performance. Second, when comparing diversity across and within chemical classes, herbivore performance was reduced when plant neighbors had greater diversity within chemical classes that are biologically inhibiting at higher quantities (i.e., quantitative defenses such as phenolics and acyl sugars). However, herbivore performance was enhanced when plant neighbors had higher levels of chemical classes that are biologically toxic (i.e., qualitative defenses such as alkaloids). Finally, herbivores performed better on plant dicultures compared to monocultures, and performance was positively associated with plant dicultures only when there were high levels of average alpha diversity within plant neighbors. Our results suggest T. ni generalist caterpillars do better when plant neighbors are chemically different because differences provide options for them to choose or to switch between plants to balance chemical uptake. Overall, herbivores interact with a large diversity of plant chemicals at multiple scales, and our results indicate that not all chemical diversity is equal: specific dimensions of phytochemical diversity have unique effects on the dynamics of herbivore performance.