Two models for AP-1 function. (A) In Model 1, AP-1 acts together with GGAs at or near the TGN, packaging a number of proteins into clathrin-coated vesicles for trafficking to peripheral endosomes. These proteins include hydrolases and their receptors, which are sorted by GGAs, with AP-1 playing an auxiliary role. AP-1 has a further role in retrieving proteins from peripheral endosomes, including empty hydrolase receptors and TGN-resident membrane proteins such as syntaxins 6, 10, and 16. Adapted from Hirst et al. (2012). (B) In Model 2, AP-1 and GGAs act independently. GGAs sort hydrolases and their receptors at or near the TGN for trafficking to endosomes, while AP-1 acts exclusively at endosomes to retrieve proteins, including empty receptors, back to the TGN. One or more of these proteins may be needed by GGAs for forward transport.