Freeze-fracture analysis of phloem structure in plant tissue cultures. I. The sieve element reticulum.

During the differentiation of phloem sieve elements, the endoplasmic reticulum undergoes unique modifications to form the sieve element reticulum (SER) which persists in mature, functioning sieve tubes. Cisternae of the SER lack ribosomes and are restricted to the periphery of the sieve element at late stages of development. Some of the SER is seen as single cisternae that are in close contact with the sieve element plasma membrane. Thin sections and freeze-fracture images of sieve elements formed in tissue cultures demonstrate that the SER consists of both single cisternae and regions of stacked cisternae at some stages of maturity. The unstacked regions of the SER are continuous with the cisternae of the stacked regions. In freeze-fracture images the single cisternae adjacent to the plasma membrane are seen to be fenestrated and the openings allow continuity between the plasma membrane and the cell lumen. It is concluded that the interface between the SER and the plasma membrane of the sieve element serves to allow membrane functions such as proton efflux, proton-sucrose cotransport and compensating movements of ions to occur in a microenvironment that is separated from the moving translocation stream in the sieve element lumen. Passage of water and translocated solutes from the plasma membrane or the SER/PM interface to the interior of the cell is enhanced by the openings in the fenestrated regions of the SER. It is suggested tha the SER may also play a role in channeling ATP from mitochondria associated with the SER to the proton-pumping ATPase in the plasma membrane and that the SER may function in the uptake and release of potassium ions in the sieve element.