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Cited by 438Cellular transport and secretionExtracellular vesicles in diseaseFungal and yeast genetics research

The exocyst complex in polarized exocytosis

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He, Bing, Guo, Wei·2009·Current Opinion in Cell Biology
DOI 10.1016/j.ceb.2009.04.007PMC PMC2725219
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Abstract

The exocyst is an octameric protein complex, which mediates the tethering of post-Golgi secretory vesicles to the plasma membrane before exocytic fusion. The exocyst assembles by side-by-side packing of rod-shaped subunits composed of helical bundles. The targeting of secretory vesicles to the plasma membrane involves direct interactions of the exocyst with PI(4,5)P(2). In addition, a number of small GTP-binding proteins interact with components of the exocyst and regulate the assembly, localization, and function of this complex. Here we review the recent advances in the field, focusing on the function of the exocyst in polarized exocytosis.

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Crossref · 10.1016/j.ceb.2009.04.007

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  • The exocyst is a protein complex that can govern the polarized cell-surface delivery of membrane and membrane proteins.

    “The exocyst is a protein complex that can govern the polarized cell-surface delivery of membrane and membrane proteins (Munson and Novick, 2006;Wu et al, 2008;He and Guo, 2009;Jin et al, 2011).”

    From Ral mediates activity‐dependent growth of postsynaptic membranes via recruitment of the exocyst (2013) · cites this paper as He and Guo, 2009

  • The subcellular localization of the exocyst in these activated muscles is consistent with the known function of the exocyst in directing post-Golgi vesicles to target membranes for localized membrane addition.

    “The subcellular localization of the exocyst in these activated muscles is consistent with the known function of the exocyst in directing post-Golgi vesicles to target membranes for localized membrane addition”

    From Ral mediates activity‐dependent growth of postsynaptic membranes via recruitment of the exocyst (2013) · cites this paper as He and Guo, 2009

  • Because the exocyst can direct membrane addition to particular membrane domains, the Ral/exocyst pathway at the NMJ likely regulates the rate and location of membrane addition, rather than other aspects of SSR development.

    “Because the exocyst can direct membrane addition to particular membrane domains (Wu et al, 2008;He and Guo, 2009), the Ral/exocyst pathway at the NMJ likely regulates the rate and location of membrane addition, rather than other aspects of SSR development.”

    From Ral mediates activity‐dependent growth of postsynaptic membranes via recruitment of the exocyst (2013) · cites this paper as He and Guo, 2009

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Research works printed but not linked: 52

  1. TerBush DR Maurice T Roth D Novick P 1996 The Exocyst is a multiprotein complex required for exocytosis in Saccharomyces cerevisiae EMBO J 15 23 6483 94 A seminal paper showing the purification of the exocyst and identification of the individual subunit in yeast 8978675 PMC452473
  2. Hsu SC Ting AE Hazuka CD Davanger S Kenny JW Kee Y Scheller RH 1996 The mammalian brain rsec6/8 complex Neuron 17 6 1209 19 First purification of mammalian exocyst complex from rat brain extract by chromatography. 8982167
  3. [3]Munson M Novick P 2006 The exocyst defrocked, a framework of rods revealed Nat Struct Mol Biol 13 7 577 81 Review 16826234
  4. Dong G Hutagalung AH Fu C Novick P Reinisch KM 2005 The structures of exocyst subunit Exo70p and the Exo84p C-terminal domains reveal a common motif Nat Struct Mol Biol 12 12 1094 100 This paper demonstrates the crystal structure of near full length of yeast Exo70 and the C-terminal domain of yeast Exo84. Exo70 and Exo84CT are principally composed of α-helices that fold into four and two helical bundles, respectively. These helical bundles pack against each other and form rod-shape structures. The authors also show that the C-terminus of Exo70 contains a number of basic residues that cluster into an electropositive surface patch 16249794
  5. [5]Hamburger ZA Hamburger AE West AP Jr Weis WI 2006 Crystal structure of the S. cerevisiae exocyst component Exo70p J Mol Biol 356 1 9 21 16359701
  6. [6]Moore BA Robinson HH Xu Z 2007 The crystal structure of mouse Exo70 reveals unique features of the mammalian exocyst J Mol Biol 371 2 410 21 17583731 PMC2692999
  7. [7]Wu S Mehta SQ Pichaud F Bellen HJ Quiocho FA 2005 Sec15 interacts with Rab11 via a novel domain and affects Rab11 localization in vivo Nat Struct Mol Biol 12 10 879 85 16155582
  8. Sivaram MV Furgason ML Brewer DN Munson M 2006 The structure of the exocyst subunit Sec6p defines a conserved architecture with diverse roles Nat Struct Mol Biol 13 6 555 6 The paper shows the crystal structure of the C-terminal domain of Sec6 composed of tandem helical-bundles. Structure comparison between Sec6CT and other exocyst components showed that they all share similar helical-bundles. These observations suggest that the exocyst subunits evolve from a common ancestor and acquire distinct surface features for their specific protein-protein interactions 16699513
  9. [9]Whyte JR Munro S 2001 The Sec34/35 Golgi transport complex is related to the exocyst, defining a family of complexes involved in multiple steps of membrane traffic Dev Cell 1 4 527 37 11703943
  10. [10]Cavanaugh LF Chen X Richardson BC Ungar D Pelczer I Rizo J Hughson FM 2007 Structural analysis of conserved oligomeric Golgi complex subunit 2 J Biol Chem 282 32 23418 26 17565980
  11. Hsu SC Hazuka CD Roth R Foletti DL Heuser J Scheller RH 1998 Subunit composition, protein interactions, and structures of the mammalian brain sec6/8 complex and septin filaments Neuron 20 6 1111 22 First structural study of the exocyst by quick-freeze/deep-etch electron microscopy 9655500
  12. Finger FP Hughes TE Novick P 1998 Sec3p is a spatial landmark for polarized secretion in budding yeast Cell 92 559 571 The first comprehensive analysis of an exocyst component in live yeast cells 9491896
  13. Boyd C Hughes T Pypaert M Novick P 2004 Vesicles carry most exocyst subunits to exocytic sites marked by the remaining two subunits, Sec3p and Exo70p J Cell Biol 167 5 889 901 An elegant fluorescence microscopic study demonstrating that different exocyst components are targeted to the plasma membrane through different mechanisms 15583031 PMC2172445
  14. [14]Zajac A Sun X Zhang J Guo W 2005 Cyclical regulation of the exocyst and cell polarity determinants for polarized cell growth Mol Biol Cell 16 1500 1512 15647373 PMC551511
  15. [15]Zhang X Zajac A Zhang J Wang P Li M Murray J Terbush D Guo W 2005 The critical role of Exo84p in the organization and polarized localization of the exocyst complex J Biol Chem 280 21 20356 20364 15788396
  16. He B Xi F Zhang X Zhang J Guo W 2007 Exo70 interacts with phospholipids and mediates the targeting of the exocyst to the plasma membrane EMBO J 26 18 4053 65 This is the first paper showing that the exocyst interacts with PI(4,5)P2 and is targeted to the plasma membrane through this interaction. The authors report that yeast Exo70 directly interact with PI(4,5)P2 through conserved basic residues located at its C-terminus. Further genetic and cell biological analyses suggest that Exo70 and Sec3 function in concert in targeting the exocyst to the plasma membrane 17717527 PMC2230670
  17. [17]Liu J Zuo X Yue P Guo W 2007 Phosphatidylinositol 4, 5-bisphosphate mediates the targeting of the exocyst to the plasma membrane for exocytosis in mammalian cells Mol Biol Cell 18 11 4483 92 17761530 PMC2043555
  18. Zhang X Orlando K He B Xi F Zhang J Zajac A Guo W 2008 Membrane association and functional regulation of Sec3 by phospholipids and Cdc42 J Cell Biol 180 1 145 58 This paper showed that Sec3 interacts with PI(4,5)P2 and disrupting the Sec3-PI(4,5)P2 or Sec3-Cdc42 interaction affects the actin-independent polarization of Sec3 in yeast 18195105 PMC2213614
  19. [19]Stroupe C Collins KM Fratti RA Wickner W 2006 Purification of active HOPS complex reveals its affinities for phosphoinositides and the SNARE Vam7p EMBO J 25 8 1579 89 16601699 PMC1440844
  20. [20]Zhang X Bi E Novick P Du L Kozminski KG Lipschutz J Guo W 2001 Cdc42 interacts with the exocyst and regulates polarized secretion J Biol Chem 276 50 46745 46750 11595741
  21. [21]Guo W Tamanoi F Novick P 2001 Spatial regulation of the exocyst complex by Rho1 GTPase Nat Cell Biol 3 4 353 360 11283608
  22. [22]Roumanie O Wu H Molk JN Rossi G Bloom K Brennwald P 2005 Rho GTPase regulation of exocytosis in yeast is independent of GTP hydrolysis and polarization of the exocyst complex J Cell Biol 170 4 583 594 16103227 PMC2171504
  23. [23]Songer JA Munson M 2009 Sec6p anchors the assembled exocyst complex at sites of secretion Mol Biol Cell 20 3 973 82 19073882 PMC2633393
  24. [24]Wedlich-Soldner R Altschuler S Wu L Li R 2003 Spontaneous cell polarization through actomyosin-based delivery of the Cdc42 GTPase Science 299 5610 1231 5 12560471
  25. [25]Grote E Carr CM Novick PJ 2000 Ordering the final events in yeast exocytosis J Cell Biol 151 2 439 52 11038189 PMC2192655
  26. [26]Sivaram MV Saporita JA Furgason ML Boettcher AJ Munson M 2005 Dimerization of the exocyst protein Sec6p and its interaction with the t-SNARE Sec9p Biochemistry 44 16 6302 11 15835919
  27. [27]Wiederkehr A De Craene JO Ferro-Novick S Novick P 2004 Functional specialization within a vesicle tethering complex: bypass of a subset of exocyst deletion mutants by Sec1p or Sec4p J Cell Biol 167 5 875 87 15583030 PMC2172455
  28. [28]Zhang X Wang P Gangar A Zhang J Brennwald P TerBush D Guo W 2005 Lethal giant larvae proteins interact with the exocyst complex and are involved in polarized exocytosis J Cell Biol 170 2 273 83 16027223 PMC2171422
  29. [29]Lehman K Rossi G Adamo JE Brennwald P 1999 Yeast homologues of tomosyn and lethal giant larvae function in exocytosis and are associated with the plasma membrane SNARE, Sec9 J Cell Biol 146 1 125 40 10402465 PMC2199738
  30. [30]Hattendorf DA Andreeva A Gangar A Brennwald PJ Weis WI 2007 Structure of the yeast polarity protein Sro7 reveals a SNARE regulatory mechanism Nature 446 7135 567 71 17392788
  31. [31]Grosshans BL Andreeva A Gangar A Niessen S Yates JR 3rd Brennwald P Novick P 2006 The yeast lgl family member Sro7p is an effector of the secretory Rab GTPase Sec4p J Cell Biol 172 1 55 66 16390997 PMC2063532
  32. [32]Bao Y Lopez JA James DE Hunziker W 2008 Snapin interacts with the Exo70 subunit of the exocyst and modulates GLUT4 trafficking J Biol Chem 283 1 324 31 17947242
  33. [33]Mima J Hickey CM Xu H Jun Y Wickner W 2008 Reconstituted membrane fusion requires regulatory lipids, SNAREs and synergistic SNARE chaperones EMBO J 27 15 2031 42 18650938 PMC2516887
  34. Novick P Field C Schekman R 1980 Identification of 23 complementation groups required for post-translational events in the yeast secretory pathway Cell 21 1 205 215 A landmark paper reporting the identification of yeast sec mutants (including six of the eight exocyst mutants) that function in the secretory pathway 6996832
  35. [35]Guo W Grant A Novick P 1999 Exo84p is an exocyst protein essential for secretion J Biol Chem 274 33 23558 64 10438536
  36. He B Xi F Zhang J TerBush D Zhang X Guo W 2007 Exo70p mediates the secretion of specific exocytic vesicles at early stages of cell cycle for polarized cell growth J Cell Biol 176 6 771 777 Demonstrated an intimate coupling between exocyst and polarized cell growth in budding yeast 17339375 PMC2064051
  37. Grindstaff KK Yeaman C Anandasabapathy N Hsu SC Rodriguez-Boulan E Scheller RH Nelson WJ 1998 Sec6/8 complex is recruited to cell-cell contacts and specifies transport vesicle delivery to the basal-lateral membrane in epithelial cells Cell 93 5 731 40 The authors show that the exocyst is recruited from cytosol to the plasma membrane at sites of early cell-cell contact during the formation of polarized epithelial monolayers. In fully polarized cells, the exocyst is enriched at the apical portion of the lateral membrane and is required for protein targeting to the basolateral domain but not the apical domain 9630218
  38. Oztan A Silvis M Weisz OA Bradbury NA Hsu SC Goldenring JR Yeaman C Apodaca G 2007 Exocyst requirement for endocytic traffic directed toward the apical and basolateral poles of polarized MDCK cells Mol Biol Cell 18 10 3978 3992 The authors carefully examined the function of the exocyst in different endocytic recycling pathways in polarized epithelial cells. The exocyst components were found to be colocalized with Rab11-positive apical recycling endosomes. Disruption of the exocyst function blocks multiple endocytic recycling pathways, including basolateral recycling, apical recycling, and basolateral-to-apical transcytosis 17686995 PMC1995710
  39. [39]Sans N Prybylowski K Petralia RS Chang K Wang YX Racca C Vicini S Wenthold RJ 2003 NMDA receptor trafficking through an interaction between PDZ proteins and the exocyst complex Nat Cell Biol 5 6 520 30 12738960
  40. Inoue M Chang L Hwang J Chiang SH Saltiel AR 2003 The exocyst complex is required for targeting of Glut4 to the plasma membrane by insulin Nature 422 629 33 This paper shows that Exo70 is a downstream effector of TC10, which promotes the cytoplasm to plasma membrane translocation of Exo70 in adipocytes in response to activation 12687004
  41. [41]Murthy M Garza D Scheller RH Schwarz TL 2003 Mutations in the exocyst component Sec5 disrupt neuronal membrane traffic, but neurotransmitter release persists Neuron 37 3 433 47 12575951
  42. Yeaman C Grindstaff KK Wright JR Nelson WJ 2001 Sec6/8 complexes on trans-Golgi network and plasma membrane regulate late stages of exocytosis in mammalian cells J Cell Biol 155 4 593 604 A comprehensive study of exocyst localization and function at the TGN and the plasma membrane 11696560 PMC2198873
  43. Prigent M Dubois T Raposo G Derrien V Tenza D Rosse C Camonis J Chavrier P 2003 ARF6 controls post-endocytic recycling through its downstream exocyst complex effector J Cell Biol 163 5 1111 21 The authors demonstrate that the exocyst is localized to the recycling endosomes and involved in recycling of an endocytosed cargo transferrin. In addition, Sec10 has been found to be a downstream effector of Arf6, which promotes Sec10 localize to plasma membrane ruffles upon activation 14662749 PMC2173613
  44. Fölsch H Pypaert M Maday S Pelletier L Mellman I 2003 The AP-1A and AP-1B clathrin adaptor complexes define biochemically and functionally distinct membrane domains J Cell Biol 163 2 351 62 This paper shows that in epithelial cells expressing AP-1B, the clathrin adaptor protein specifically expressed in epithelial cells and mediating the trafficking of proteins to the basolateral membrane, promotes the recruitment of the exocyst to the recycling endosomes 14581457 PMC2173537
  45. [45]Zhang XM Ellis S Sriratana A Mitchell CA Rowe T 2004 Sec15 is an effector for the Rab11 GTPase in mammalian cells J Biol Chem 279 41 43027 34 15292201
  46. Guo W Roth D Walch-Solimena C Novick P 1999 The exocyst is an effecter for Sec4p, targeting secretory vesicles to sites of exocytosis EMBO J 18 1071 1080 This is the first paper showing that the exocyst is a downstream effector of the Ras family of small GTPase. The authors demonstrate that yeast Sec15 directly interacts with GTP-bound form of Rab GTPase Sec4 but not other Rabs; mutations in Sec4 inhibited exocyst assembly. This study suggests a mechanism by which the Rab GTPases controls specificity of different vesicular trafficking events by regulating specific downstream vesicle tethering factors 10022848 PMC1171198
  47. Langevin J Morgan MJ Sibarita JB Aresta S Murthy M Schwarz T Camonis J Bellaiche Y 2005 Drosophila exocyst components Sec5, Sec6, and Sec15 regulate DE-Cadherin trafficking from recycling endosomes to the plasma membrane Dev Cell 9 3 355 76 This paper shows that loss of function of Sec5, Sec6, or Sec15 in Drosophila epithelial cells inhibits recycling of E-cadherin to the plasma membrane and results in its accumulation in enlarged Rab11-positive recycling endosomes 16224820
  48. [48]Adamo JE Rossi G Brennwald P 1999 The Rho GTPase Rho3 has a direct role in exocytosis that is distinct from its role in actin polarity Mol Biol Cell 10 4121 4133 10588647 PMC25747
  49. [49]Robinson NG Guo L Imai J Toh EA Matsui Y Tamanoi F 1999 Rho3 of Saccharomyces cerevisiae, which regulates the actin cytoskeleton and exocytosis, is a GTPase which interacts with Myo2 and Exo70 Mol Cell Biol 19 5 3580 3587 10207081 PMC84150
  50. [50]Hutagalung AH Coleman J Pypaert M Novick PJ 2009 An internal domain of Exo70p is required for actin-independent localization and mediates assembly of specific exocyst components Mol Biol Cell 20 1 153 63 18946089 PMC2613103
  51. Adamo JE Moskow JJ Gladfelter AS Viterbo D Lew DJ Brennwald PJ 2001 Yeast Cdc42 functions at a late step in exocytosis, specifically during polarized growth of the emerging bud J Cell Biol 155 4 581 592 Demonstrated a role of Cdc42 in kinetic regulation of exocytosis during yeast budding 11706050 PMC2198861
  52. Moskalenko S Henry DO Rosse C Mirey G Camonis JH White MA 2002 The exocyst is a Ral effector complex Nat Cell Biol 4 1 66 72 Together with other labs, demonstrated that the exocyst is a direct downstream effector of Ral in mammalian cells 11740492

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Authors
He, B., & Guo, W.
Affiliations
University of Pennsylvania (US)
Corresponding author
Guo, Wei
ORCID
Venue
Current Opinion in Cell Biology
Publisher
Elsevier BV
Published
2009-08
Volume
21
Issue
4
Pages
537-542
ISSN
0955-0674 (ISSN-L)
DOI
10.1016/j.ceb.2009.04.007
PMC
PMC2725219
OpenAlex
W2109662154
Cited by
438
References
0
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FWCI
3.86
Type
journal article
Language
en
Open access
Licence
Locations
  • Current Opinion in Cell Biology (publishedVersion)
  • PubMed (publishedVersion)
  • Europe PMC (PubMed Central) (acceptedVersion)
  • PubMed Central (acceptedVersion)
Funders
  • National Institutes of Health
  • National Institute of General Medical Sciences (R01GM085146, R01GM064690)
Topics
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Cell Biology › Cellular transport and secretion
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Extracellular vesicles in disease
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Fungal and yeast genetics research
Keywords
exocyst complex, polarized exocytosis, post-Golgi vesicles, plasma membrane, phosphatidylinositol 4,5-bisphosphate, small GTPases, membrane trafficking
MeSH terms
Animals, Cell Membrane, Exocytosis, Golgi Apparatus, Guanosine Triphosphate, HeLa Cells, Humans, Kinetics, Models, Biological, Protein Binding, Protein Structure, Tertiary, GTP-Binding Proteins, SNARE Proteins

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