Nucleus-encoded Precursors to Thylakoid Lumen Proteins of Euglena gracilis Possess Tripartite Presequences

The complete presequences of the nucleus-encoded precursors to two proteins, cytochrome c6 and the 30-kDa protein of the oxygen-evolving complex, that reside in the thylakoid lumen of the chloroplasts of Euglena gracilis are presented. Sorting of these proteins involves translocation across four membranes, the three-membraned chloroplast envelope and the thylakoid membrane. The tripartite presequences show the structure: signal sequence transit sequence signal sequence. Three hydrophobic domains become apparent: two of them correspond to signal sequences for translocation across the endoplasmic reticulum (ER) membrane and the thylakoid membrane, respectively, whereas the third constitutes the stop-transfer signal contained in the long stroma-targeting part of the tripartite presequence.