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Volatiles are often released upon herbivory as plant defense compounds. While the formation of volatiles above-ground has been intensively studied, little is known about herbivore-induced root volatiles. Here, we show that cockchafer larvae-damaged roots of  Populus trichocarpa  and  P. nigra  release a mixture of monoterpenes, including (−)-α-pinene, (−)-camphene, (−)-β-pinene,  p -cymene, and 1,8-cineole. Three terpene synthases, PtTPS16 and PtTPS21 from  P. trichocarpa  and PnTPS4 from  P. nigra , could be identified and characterized  in vitro . PnTPS4 was found to produce 1,8-cineole as sole product. PtTPS16 and PtTPS21, although highly similar to each other, showed different product specificities and produced γ-terpinene and a mixture of (−)-camphene, (−)-α-pinene, (−)-β-pinene, and (−)-limonene, respectively. Four active site residues were found to determine the different product specificities of the two enzymes. The expression profiles of  PtTPS16, PtTPS 2 1 , and  PnTPS4  in undamaged and herbivore-damaged poplar roots generally matched the emission pattern of monoterpenes, indicating that monoterpene emission in roots is mainly determined at the gene transcript level. Bioassays with  Phytophtora cactorum  (Oomycetes) revealed inhibitory effects of vapor-phase 1,8-cineole and (−)-β-pinene on the growth of this important plant pathogen. Thus herbivore-induced volatile monoterpenes may have a role in defense against pathogens that cause secondary infections after root wounding.

The occurrence and formation of monoterpenes in herbivore-damaged poplar roots