Redefining the metabolic continuity of chloroplasts and ER.

As a hub for plant metabolism, plastids extensively exchange metabolites with the extraplastid environment. For polar metabolites, membrane transporters mediate this exchange, but for many plastid-synthesized nonpolar compounds, such transporters remain elusive. Here, we discuss recent data from transorganellar complementation studies that demonstrate that enzymes in one organelle can directly access nonpolar metabolites from a companion organelle. We propose that a mechanism, based on hemifused-membranes at plastid-endoplasmic reticulum (ER) contact sites, facilitates interorganellar interactions and allows enzymes direct, transporter-independent access to a range of nonpolar compounds in both organelle membranes. In a wider context, interorganellar metabolism at hemifusion interfaces would allow evolution of membrane-spanning pathways for the many thousands of nonpolar metabolites in the plant kingdom to be uncoupled from coevolution with nonpolar metabolite transporters.