Shoot–root defense signaling and activation of root defense by leaf damage in poplar1

Shoot–root systemic defense signaling of hybrid poplar (Populus trichocarpa Torr. & A. Gray Populus deltoides Bartr. ex Marsh.) was investigated with molecular techniques to extend existing knowledge of poplar defense. Treatment of roots with methyl jasmonate demonstrated that transcripts of PtdTI3, a poplar trypsin inhibitor and marker of poplar defense responses, can be induced in poplar roots as well as leaves. Moreover, simulated herbivory of poplar leaves with methyl jasmonate treatment or wounding with pliers also induced PtdTI3 mRNA in roots, which implies downward, or basipetal, systemic signaling from shoots to roots. In addition, the inducible root-defense response comprised both increased PtdTI3 protein levels and trypsin-inhibitor activity. The inducible systemic response was further investigated with comparative macroarray analyses which indicated that in addition to PtdTI3, other genes respond in roots after wounding and methyl jasmonate treatment of leaves. The majority of the 17 genes encode previously identified leaf herbivory defense genes; however, some genes strongly up-regulated in leaves were not induced in roots. The identification of multiple defense genes that are inducible in roots following leaf damage is clear evidence of a systemic defense response in roots and the presence of basipetal shoot–root defense signaling.