Transport - Clathrin-coated vesicle cycle

Clathrin-coated vesicle cycle

Transport vesicles are classified according to the components of the protein coat that surrounds them during their genesis and early life. One of the most common and probably best-characterized classes of coated vesicle is that comprising three-layered Clathrin-coated vesicles (CCVs). CCVs are so-called because the main component of the coat is complex Clathrin, which forms a polymeric mechanical scaffold on the vesicle surface. The inner, membrane layer with its embedded cargo is linked to the outer, Clathrin layer by a middle layer that consists of various clathrin-adaptor molecules and other proteins that have accessory/regulatory roles in CCV assembly. The classical key components of absolute majority cargos Clathrin-dependent endocytosis are AP complex 2, Eps15, AP180, Epsin 1, HIP1/ HIP12 and others [1], [2], [3].

After the Clathrin lattice is formed, dynamins (e.g., Dynamin-2), endophilin (e.g., Endophilin B1), epsins and amphiphysin (e.g., BIN1 (Amphiphysin II)) are involved in membrane invagination and Clathrin rearrangements. The plus-end motor Myosin I pulls the Dynamin-2 ring in the direction of the cell surface, while the minus-end motor Myosin VI pulls the coated bud into the cytosol. The resulting strain could then sever the constricted stalk beneath the dynamin ring [4].

Next phase is a fusion of coated-pit-derived primary endocytic vesicles with sorting endosomes. It is regulated by member RAS oncogene family Rab-5A, EEA1 [5], [6], [7] and Soluble N-ethylmaleimide-sensitive factor attachment protein receptors (SNARE) [7], [8].

The maturation of sorting endosomes to late endosomes is realized with participation member RAS oncogene family Rab-7 via unknown mechanism [9], [10].

Late endosomes may participate in fusion to other late endosomes or lysosome via SNARE-mediated mechanism. Combinatorial SNARE complexes with VAMP7 or VAMP8 define these different late endocytic fusion events, accordingly [8], [11], [12].

In addition, Rab-7 is directly involved in the aggregation and fusion of late endocytic structures/lysosomes [11], [13].

Then, cargo may be delivered to the Golgi from late endosomes with participation RAS oncogene family Rab-9 [10], [14], [15]. In addition, proteins from endoplasmic reticulum may be translocated to the COPII-dependent manner [16].

Modified in endoplasmic reticulum and/or Golgi cargo may be delivered from the Golgi back to the cell surface, possibly with participation Rab-8/ Optineurin/ Myosin VI pathway [7], [17], [18] and/or coat protein complex termed Coatomer [7], [19], [20].

Moreover, cargo may be delivered to the cell surface via shot pathway from endosomes via different recycling endosomes [7]. It is realized mainly via Rab-4 and/or Rab-11A-dependent mechanisms [10], [21], [22].

It is shown, that recycling endosomes (endocytic recycling) to Golgi traffic may be realized via different SNARE complexes [7], [8], [23].

Objects list:

References:

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