Kd. Klepzig et al., EFFECTS OF BIOTIC AND ABIOTIC STRESS ON INDUCED ACCUMULATION OF TERPENES AND PHENOLICS IN RED PINES INOCULATED WITH BARK BEETLE-VECTORED FUNGUS, Journal of chemical ecology, 21(5), 1995, pp. 601-626
This study characterized the chemical response of healthy red pine to
artificial inoculation with the bark beetle-vectored fungus Leptograph
ium terebrantis. In addition, we sought to determine whether stress al
tered this induced response and to understand the implications of thes
e interactions to the study of decline diseases. Twenty-five-year-old
trees responded to mechanical wounding or inoculation with L. terebran
tis by producing resinous reaction lesions in the phloem. Aseptically
wounded and wound-inoculated phloem contained higher concentrations of
phenolics than did constitutive tissue. Trees inoculated with L. tere
brantis also contained higher concentrations of six monoterpenes, alph
a-pinene, beta-pinene, 3-carene, limonene, camphene, and myrcene, and
higher total monoterpenes than did trees that were mechanically wounde
d or left unwounded. Concentrations of these moneterpenes increased wi
th time after inoculation. Total phenolic concentrations in unwounded
stem tissue did not differ between healthy and root-diseased trees. Li
kewise, constitutive monoterpene concentrations in stem phloem were si
milar between healthy and root-diseased trees. However, when stem phlo
em tissue was challenged with fungal inoculations, reaction tissue fro
m root-diseased trees contained lower concentrations of alpha-pinene,
the predominant monoterpene in red pine, than did reaction tissue from
healthy trees. Seedlings stressed by exposure to low light levels exh
ibited less extensive induced chemical changes when challenge inoculat
ed with L. terebrantis than did seedlings growing under higher light.
Stem phloem tissue in these seedlings contained lower concentrations o
f ol-pinene than did nonstressed seedlings also challenge inoculated w
ith L. terebrantis. It is hypothesized that monoterpenes and phenolics
play a role in the defensive response of red pine against insect-fung
al attack, that stress may predispose red pine to attack by insect-fun
gal complexes, and that such interactions are involved in red pine dec
line disease. Implications to plant defense theory and interactions am
ong multiple stress agents in forest decline are discussed.