• Eukaryotic cells contain many membrane-enclosed organelles, including a nucleus, an endoplasmic reticulum (ER), a Golgi apparatus, lysosomes, endosomes, mitochondria, chloroplasts (in plant
cells), and peroxisomes. The ER, Golgi apparatus, peroxisomes, endosomes, and lysosomes are all part of the endomembrane system.
- Most organelle proteins are made in the cytosol and transported into the organelle where they function. Sorting signals in the amino acid sequence guide the proteins to the correct organelle; proteins that function in the cytosol have no such signals and remain where they are made.
- Nuclear proteins contain nuclear localization signals that help direct their active transport from the cytosol into the nucleus through nuclear pores, which penetrate the double membrane of the nuclear envelope. The proteins are transported in their fully folded conformation.
- Most mitochondrial and chloroplast proteins are made in the cytosol and are then transported into the organelles by protein translocators in their membranes. The proteins are unfolded during the transport process.
- The ER makes most of the cell’s lipids and many of its proteins. The proteins are made by ribosomes that are directed to the ER by a signal-recognition particle (SRP) in the cytosol that recognizes an ER signal sequence on the growing polypeptide chain. The ribosome–SRP complex binds to a receptor on the ER membrane, which passes the ribosome to a protein translocator that threads the growing polypeptide across the ER membrane.
• Water-soluble proteins destined for secretion or for the lumen of an organelle of the endomembrane system pass completely into the ER lumen,
while transmembrane
proteins destined for either the membrane of these organelles or for the plasma membrane remain
anchored in the lipid bilayer by one or more
membrane-spanning α helices.
• In the ER lumen, proteins fold up, assemble
with their protein partners, form disulfide bonds, and become decorated with oligosaccharide
chains.
• Exit from the ER is an important quality-control step; proteins that either fail to fold properly or
fail to assemble with their normal partners are
retained in the ER by chaperone proteins, which
prevent heir aggregation and help them fold;
proteins that still fail to fold or assemble are
transported to the cytosol, where they are
degraded.
• Excessive accumulation of misfolded proteins
triggers an unfolded protein response that expands the ER, increases its capacity to fold new proteins properly, and reduces protein synthesis.
• Protein transport from the ER to the Golgi
apparatus and from the Golgi apparatus to other
destinations is mediated by transport vesicles that continually bud off from one membrane and fuse with another, a process called vesicular transport.
• Budding transport vesicles have distinctive coat proteins on their cytosolic surface; the assembly
of the coat helps drive both the budding process
and the incorporation of cargo receptors, with
their bound cargo molecules, into the forming
vesicle.
- Coated vesicles rapidly lose their protein coat, enabling them to dock and then fuse with a particular target membrane; docking and fusion are mediated by proteins on the surface of the vesicle and target membrane, including Rab, tethering, and SNARE proteins.
- The Golgi apparatus receives newly made proteins from the ER; it modifies their oligosaccharides, sorts the proteins, and dispatches them from the trans Golgi network to the plasma membrane, lysosomes (via endosomes), or secretory vesicles.
- In all eukaryotic cells, transport vesicles continually bud from the trans Golgi network and fuse with the plasma membrane; this process of constitutive exocytosis delivers proteins to the cell surface for secretion and incorporates lipids and proteins into the plasma membrane.
- Specialized secretory cells also have a regulated exocytosis pathway, in which molecules concentrated and stored in secretory vesicles are released from the cell by exocytosis when the cell is signaled to secrete.
• Cells ingest fluid, molecules, and sometimes
even particles by endocytosis, in which regions of plasma membrane invaginate and pinch off to
form endocytic vesicles.
• Much of the material that is endocytosed is
delivered to endosomes, which mature into lysosomes, in which the material is degraded by
hydrolytic enzymes; most of the components of
the endocytic vesicle membrane, however, are
recycled in transport vesicles back to the plasma membrane for
reuse.