Bracelet salt glands of the recretohalophyte Limonium bicolor:Distribution,morphology,and induction

Halophytes complete their life cycles in saline environments.The recretohalophyte Limonium bicolor has evolved a specialized salt secretory structure,the salt gland,which excretes Na+to avoid salt damage.Typical L.bicolor salt glands consist of 16 cells with four fluorescent foci and four secretory pores.Here,we describe a special type of salt gland at the base of the L.bicolor leaf petiole named bracelet salt glands due to their beaded-bracelet-like shape of blue auto-fluorescence.Bracelet salt glands contain more than 16 cells and more than four secretory pores.Leaf disc secretion measurements and non-invasive micro-test techniques indicated that bracelet salt glands secrete more salt than normal salt glands,which helps maintain low Na+levels at the leaf blade to protect the leaf.Cytokinin treatment induced bracelet salt gland differentiation,and the developed ones showed no further differentiation when traced with a living fluorescence microscopy imager,even though new salt gland development and leaf expansion were observed.Transcriptome revealed a NAC transcription factor gene that participates in bracelet salt gland development,as confirmed by its genome editing and overexpression in L.bicolor.These findings shed light on bracelet salt gland development and may facilitate the engineering of salt-tolerant crops.