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Access Type

Open Access Thesis

Document Type


Degree Program

Organismic & Evolutionary Biology

Degree Type

Master of Science (M.S.)

Year Degree Awarded


Month Degree Awarded



Eastern hemlock hosts the hemlock woolly adelgid, an introduced sap-feeding insect that causes rapid deterioration of the host. Like most conifers, eastern hemlock produces a variety of constitutive and induced defenses, primarily terpenoids. To explore the relationship of terpenoid defenses with adelgid infestations, we artificially infested hemlocks at a forest site and a plantation site, and compared their terpenoid concentrations to those in control trees. Infested trees showed lower terpenoid concentrations than control trees, suggesting that eastern hemlock not only fails to induce production of terpenoids in response to adelgid infestation, but becomes less able to produce carbon-based defenses due to loss of carbon resources to the adelgid. Greater light intensity may account for consistently higher terpenoid concentrations at the plantation site, supporting the explanation that carbon limitation restricts terpenoid production.

Recent studies have identified a small number of individual eastern hemlock trees that demonstrate relative resistance to the hemlock woolly adelgid. We compared concentrations of terpenoids in susceptible and relatively resistant trees, both in the forest and in propagated cuttings in a common-garden setting. Terpenoid concentrations were higher in twig tissue of resistant versus susceptible trees, across six sampling dates and at both sites. Because the common-garden cuttings were free of herbivores, the higher terpenoid concentrations are interpreted as a constitutive defense. Increased levels of monoterpenes and sesquiterpenes imply an overall increase in the input of carbon precursors to both terpenoid synthesis pathways. This result suggests either an altered growth-defense balance favoring allocation of carbon resources towards production of defenses, or overall greater carbon availability in growing twig tissue of adelgid-resistant eastern hemlock individuals.

We contribute detailed terpenoid data to the study of the eastern hemlock – hemlock woolly adelgid system. Our solvent extraction method permits us to examine needle and twig tissues separately, capture minor components at low concentrations, and focus on stored rather than volatilized terpenoids. By relating terpenoid concentrations to insect densities, we explore the relationships of tentatively defensive chemistry to insect population dynamics. The question remains which terpenoids, if any, directly affect hemlock woolly adelgid and what role phenols may play in the system.


First Advisor

Joseph S Elkinton