ERES (ER exit sites) and the "secretory unit concept"

The higher plant Golgi apparatus consists of hundreds of individual Golgi stacks which move along the cortical ER, propelled by the actomysin system. Anterograde and retrograde transport between the endoplasmic reticulum (ER) and the plant Golgi occurs over a narrow interface (around 500 nm) and is generally considered to be mediated by COP-coated vesicles. Previously, ER exit sites (ERES) have been identified on the basis of to localization of transiently expressed COPII-coat proteins. As a consequence it has been held that ERES in higher plants are intimately associated with Golgi stacks, and that both move together as an integrated structure: the "secretory unit". Using a new COPII marker, as well as YFP-SEC24 (a bona fide COPII coat protein), we have made observations on tobacco leaf epidermis at high resolution in the CLSM. Our data clearly shows that COPII fluorescence is associated with the Golgi stacks rather than the surface of the ER and probably represents the temporary accumulation of COPII vesicles in the Golgi matrix prior to fusion with the cis-Golgi cisternae. We have calculated the numbers of COPII vesicles which would be required to provide a typical Golgi-associated COPII-fluorescent signal as being much less than 20. We have discussed the consequences of this and question the continued usage of the term "secretory unit".