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During phloem unloading, multiple cell-to-cell transport events move organicsubstances to the root meristem. Although the primary unloading event from thesieve elements to the phloem pole pericycle has been characterized to someextent, little is known about post-sieve element unloading. Here, we report anovel gene, PHLOEM UNLOADING MODULATOR (PLM), in the absence of whichplasmodesmata-mediated symplastic transport through the phloem polepericycle--endodermis interface is specifically enhanced. Increased unloadingis attributable to a defect in the formation of the endoplasmicreticulum--plasma membrane tethers during plasmodesmal morphogenesis, resultingin the majority of pores lacking a visible cytoplasmic sleeve. PLM encodes aputative enzyme required for the biosynthesis of sphingolipids withvery-long-chain fatty acid. Taken together, our results indicate thatpost-sieve element unloading involves sphingolipid metabolism, which affectsplasmodesmal ultrastructure. They also raise the question of how and whyplasmodesmata with no cytoplasmic sleeve facilitate molecular trafficking.

Sphingolipid biosynthesis modulates plasmodesmal ultrastructure and phloem unloading