Importin 1 Mediates Nuclear Entry of EIN2C to Confer the Phloem-Based Defense against Aphids

Ethylene Insensitive 2 (EIN2) is an integral membrane protein that regulates ethylene signaling towards plant development and immunity by release of its carboxy-terminal functional portion (EIN2C) into the nucleus. The present study elucidates that the nuclear trafficking of EIN2C is induced by importin beta 1, which triggers the phloem-based defense (PBD) against aphid infestations in Arabidopsis. In plants, IMP beta 1 interacts with EIN2C to facilitate EIN2C trafficking into the nucleus, either by ethylene treatment or by green peach aphid infestation, to confer EIN2-dependent PBD responses, which, in turn, impede the phloem-feeding activity and massive infestation by the aphid. In Arabidopsis, moreover, constitutively expressed EIN2C can complement the imp b1 mutant regarding EIN2C localization to the plant nucleus and the subsequent PBD development in the concomitant presence of IMP beta 1 and ethylene. As a result, the phloem-feeding activity and massive infestation by green peach aphid were highly inhibited, indicating the potential value of EIN2C in protecting plants from insect attacks.