Plasmodesmata in phloem: different gateways for different cargoes

The long-distance transport of sugars and nutrients through the phloem is essential for the proper function and growth of vascular plants. However, in addition to essential nutrients and sugars, phloem sap also contains small molecules (e.g. hormones) as well as a diverse population of macromolecules (i.e. proteins small RNAs, and mRNAs), the endogenous functions of which remain largely unknown. Understanding the cellular origins of these mobile macromolecules, their path into and out of the phloem translocation stream, and their fate at their new destination is essential for characterizing their presumptive function. Specialized plasmodesmal connections that regulate phloem entry and exit are central to all of these processes. Here, we highlight new discoveries underscoring plasmodesmal structure and function during unloading of various molecules in the sink, and discuss how these findings shape a new view for the potential function of phloem-mobile macromolecules.