Tomato ABA-IMPORTING TRANSPORTER 1.1 inhibits seed germination under high salinity conditions

The plant hormone abscisic acid (ABA) plays a central role in the regulation of seed maturation and dormancy. ABA also restrains germination under abiotic-stress conditions. Here, we show in tomato (Solanum lycopersicum) that the ABA importer ABA-IMPORTING TRANSPORTER 1.1 (AIT1.1/NPF4.6) has a role in radicle emergence under salinity conditions. AIT1.1 expression was upregulated following seed imbibition, and CRISPR/Cas9-derived ait1.1 mutants exhibited faster radicle emergence, increased germination and partial resistance to ABA. AIT1.1 was highly expressed in the endosperm, but not in the embryo, and ait1.1 isolated embryos did not show resistance to ABA. On the other hand, loss of AIT1.1 activity promoted the expression of endosperm-weakening-related genes, and seed-coat scarification eliminated the promoting effect of ait1.1 on radicle emergence. Therefore, we propose that imbibition-induced AIT1.1 expression in the micropylar endosperm mediates ABA-uptake into micropylar cells to restrain endosperm weakening. While salinity conditions strongly inhibited wild-type M82 seed germination, high salinity had a much weaker effect on ait1.1 germination. We suggest that AIT1.1 evolved to inhibit germination under unfavorable conditions, such as salinity. Unlike other ABA mutants, ait1.1 exhibited normal seed longevity, and therefore, the ait1.1 allele may be exploited to improve seed germination in crops.