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Aims  To elucidate the genetics underlying salt tolerance in recretohalophytes and assess its relevance to non-halophytes, we cloned the  Limonium bicolor  homolog of  Arabidopsis thaliana  (Arabidopsis)  SUPER SENSITIVE TO ABA AND DROUGHT2  ( AtSAD2 ) and named it  LbSAD2 , an importin-β gene associated with trichome initiation and reduced abscisic acid (ABA) sensitivity, and then we assessed the heterologously expressed  LbSAD2  in Arabidopsis.    Methods  We examined  LbSAD2  expression and assessed the effect of heterologous  LbSAD2  expression in Arabidopsis on root hair/trichome induction; the expression levels of possible related genes in trichome/root hair development; some physiological parameters involved in salt tolerance including germination rate, root length, and contents of Na + , proline, and malondialdehyde; and the response of ABA at the germination stage.    Results  The  LbSAD2  gene is highly expressed in the salt gland development stage and salt treatment, especially located in the salt gland by  in situ  hybridization, and the LbSAD2 protein contains some special domains compared with AtSAD2, which may suggest the involvement of LbSAD2 in salt tolerance. Compared with the  SAD2/GL1  mutant CS65878, which lacks trichomes,  CS65878-35S:LbSAD2  had higher trichome abundance but lower root hair abundance. Under 100 mM NaCl treatment,  CS65878-35S:LbSAD2  showed enhanced germination and root lengths; improved physiological parameters, including high proline and low contents of Na +  and malondialdehyde; higher expression of the salt-tolerance genes Δ 1-PYRROLINE-5-CARBOXYLATE SYNTHETASE 1  ( P5CS1 ) and  GST CLASS TAU 5  ( GSTU5 ); reduced ABA sensitivity; and increased expression of the ABA signaling genes  RESPONSIVE TO ABA 18  ( RAB18 ) and  SNF1-RELATED PROTEIN KINASE 2  ( SRK2E ), but not of the ABA biosynthesis gene  9-CIS-EPOXYCAROTENOID DIOXYGENASE 3  ( NCED3 ).    Conclusion   LbSAD2  enhances salt tolerance in Arabidopsis by specifically reducing root hair development, Na +  accumulation, and ABA sensitivity.

frontiers in plant science

Importin-β From the Recretohalophyte Limonium bicolor Enhances Salt Tolerance in Arabidopsis thaliana by Reducing Root Hair Development and Abscisic Acid Sensitivity