Terpenes Tell Different Tales at Different Scales: Glimpses into the Chemical Ecology of Conifer - Bark Beetle - Microbial Interactions

Chemical signaling mediates nearly all aspects of species interactions. Our knowledge of these signals has progressed dramatically, and now includes good characterizations of the bioactivities, modes of action, biosynthesis, and genetic programming of numerous compounds affecting a wide range of species. A major challenge now is to integrate this information so as to better understand actual selective pressures under natural conditions, make meaningful predictions about how organisms and ecosystems will respond to a changing environment, and provide useful guidance to managers who must contend with difficult trade-offs among competing socioeconomic values. One approach is to place stronger emphasis on cross-scale interactions, an understanding of which can help us better connect pattern with process, and improve our ability to make mechanistically grounded predictions over large areas and time frames. The opportunity to achieve such progress has been heightened by the rapid development of new scientific and technological tools. There are significant difficulties, however: Attempts to extend arrays of lower-scale processes into higher scale functioning can generate overly diffuse patterns. Conversely, attempts to infer process from pattern can miss critically important lower-scale drivers in systems where their biological and statistical significance is negated after critical thresholds are breached. Chemical signaling in bark beetle - conifer interactions has been explored for several decades, including by the two pioneers after whom this award is named. The strong knowledge base developed by many researchers, the importance of bark beetles in ecosystem functioning, and the socioeconomic challenges they pose, establish these insects as an ideal model for studying chemical signaling within a cross-scale context. This report describes our recent work at three levels of scale: interactions of bacteria with host plant compounds and symbiotic fungi (tree level, biochemical time), relationships among inducible and constitutive defenses, population dynamics, and plastic host-selection behavior (stand level, ecological time), and climate-driven range expansion of a native eruptive species into semi-na < ve and potentially na < ve habitats (geographical level, evolutionary time). I approach this problem by focusing primarily on one chemical group, terpenes, by emphasizing the curvilinear and threshold-structured basis of most underlying relationships, and by focusing on the system's feedback structure, which can either buffer or amplify relationships across scales.