Phloem development: current knowledge and future perspectives.

Phloem, as a major tissue mediating long-distance communication, has been an object of extensive research ever since its structure was first reported in 1837. Functional phloem consists of sieve elements (SEs) and companion cells (CCs). While SEs are enucleated conducting cells in the phloem, CCs are cells with intact cellular components and are known to support the functioning of SEs. CCs are closely linked to SEs by symplastic connections mediated by plasmodesmata (PD). Sieve elements are notoriously sensitive to manipulation, which has hampered efforts to investigate their structure using microscopy or histology; phloem thus remains a mysterious tissue almost 200 yr after its discovery. Nevertheless, consistent efforts have overcome many of the technical barriers and generated considerable amounts of data about the structure and function of phloem. Advances in the 1950s and 1960s significantly improved our understanding of phloem anatomy and function. A major function of the phloem is to establish symplastic connections throughout the plant body, delivering nutrients and various signaling molecules, which play pivotal roles in growth and development. Despite the importance of phloem, details about the molecular mechanisms responsible for the establishment and maintenance of phloem continuity remain elusive.