1. Spooner RA, Smith DC, Easton AJ, Roberts LM, Lord JM (2006) Retrograde
transport pathways utilised by viruses and protein toxins. Virol J 3: 26.
2. Bellisola G, Fracasso G, Ippoliti R, Menestrina G, Rosen A, et al. (2004)
Reductive activation of ricin and ricin A-chain immunotoxins by protein
disulfide isomerase and thioredoxin reductase. Biochem Pharmacol 67:
1721–1731.
3. Spooner RA, Watson PD, Marsden CJ, Smith DC, Moore KA, et al. (2004)
Protein disulphide-isomerase reduces ricin to its A and B chains in the
endoplasmic reticulum. Biochem J 383: 285–293.
4. Mayerhofer PU, Cook JP, Wahlman J, Pinheiro TT, Moore KA, et al. (2009)
Ricin A chain insertion into endoplasmic reticulum membranes is triggered by a
temperature increase to 37uC. J Biol Chem 284: 10232–10242.
5. Ye Y, Shibata Y, Yun C, Ron D, Rapoport TA (2004) A membrane protein
complex mediates retro-translocation from the ER lumen into the cytosol.
Nature 429: 841–847.
6. Sun F, Zhang R, Gong X, Geng X, Drain PF, et al. (2006) Derlin-1 promotes
the efficient degradation of the cystic fibrosis transmembrane conductance
regulator (CFTR) and CFTR folding mutants. J Biol Chem 281: 36856–36863.
7. Tsai YC, Weissman AM (2010) The Unfolded Protein Response, Degradation
from Endoplasmic Reticulum and Cancer. Genes Cancer 1: 764–778.
8. Rabinovich E, Kerem A, Frohlich KU, Diamant N, Bar-Nun S (2002) AAAATPase
p97/Cdc48p, a cytosolic chaperone required for endoplasmic
reticulum-associated protein degradation. Mol Cell Biol 22: 626–634.
9. Elkabetz Y, Shapira I, Rabinovich E, Bar-Nun S (2004) Distinct steps in
dislocation of luminal endoplasmic reticulum-associated degradation substrates:
roles of endoplamic reticulum-bound p97/Cdc48p and proteasome. J Biol
Chem 279: 3980–3989.
10. Ye Y, Meyer HH, Rapoport TA (2001) The AAA ATPase Cdc48/p97 and its
partners transport proteins from the ER into the cytosol. Nature 414: 652–656.
11. Stirling CJ, Lord JM (2006) Quality control: linking retrotranslocation and
degradation. Curr Biol 16: R1035–1037.
12. Wang Q, Li L, Ye Y (2006) Regulation of retrotranslocation by p97-associated
deubiquitinating enzyme ataxin-3. J Cell Biol 174: 963–971.
13. Zhong X, Pittman RN (2006) Ataxin-3 binds VCP/p97 and regulates
retrotranslocation of ERAD substrates. Hum Mol Genet 15: 2409–2420.
14. Medicherla B, Kostova Z, Schaefer A, Wolf DH (2004) A genomic screen
identifies Dsk2p and Rad23p as essential components of ER-associated
degradation. EMBO Rep 5: 692–697.
15. Wesche J, Rapak A, Olsnes S (1999) Dependence of ricin toxicity on
translocation of the toxin A-chain from the endoplasmic reticulum to the
cytosol. J Biol Chem 274: 34443–34449.
16. Willer M, Forte GM, Stirling CJ (2008) Sec61p is required for ERAD-L: genetic
dissection of the translocation and ERAD-L functions of Sec61P using novel
derivatives of CPY. J Biol Chem 283: 33883–33888.
17. Slominska-Wojewodzka M, Gregers TF, Walchli S, Sandvig K (2006) EDEM is
involved in retrotranslocation of ricin from the endoplasmic reticulum to the
cytosol. Mol Biol Cell 17: 1664–1675.
18. Redmann V, Oresic K, Tortorella LL, Cook JP, Lord M, et al. (2011)
Dislocation of ricin toxin A chains in human cells utilizes selective cellular
factors. J Biol Chem 286: 21231–21238.
19. Spooner RA, Hart PJ, Cook JP, Pietroni P, Rogon C, et al. (2008) Cytosolic
chaperones influence the fate of a toxin dislocated from the endoplasmic
reticulum. Proc Natl Acad Sci U S A 105: 17408–17413.
20. Endo Y, Tsurugi K (1987) RNA N-glycosidase activity of ricin A-chain.
Mechanism of action of the toxic lectin ricin on eukaryotic ribosomes. J Biol
Chem 262: 8128–8130.
21. Li S, Spooner RA, Allen SC, Guise CP, Ladds G, et al. (2010) Foldingcompetent
and folding-defective forms of ricin A chain have different fates after
retrotranslocation from the endoplasmic reticulum. Mol Biol Cell 21:
2543–2554.
22. Endo Y, Tsurugi K, Yutsudo T, Takeda Y, Ogasawara T, et al. (1988) Site of
action of a Vero toxin (VT2) from Escherichia coli O157:H7 and of Shiga toxin
on eukaryotic ribosomes. RNA N-glycosidase activity of the toxins. Eur J Biochem
171: 45–50.
23. Yu M, Haslam DB (2005) Shiga toxin is transported from the endoplasmic
reticulum following interaction with the luminal chaperone HEDJ/ERdj3. Infect
Immun 73: 2524–2532.
24. Falguieres T, Johannes L (2006) Shiga toxin B-subunit binds to the chaperone
BiP and the nucleolar protein B23. Biol Cell 98: 125–134.
25. Williams DB, Watts TH (1995) Molecular chaperones in antigen presentation.
Curr Opin Immunol 7: 77–84.
26. Cresswell P (2000) Intracellular surveillance: controlling the assembly of MHC
class I-peptide complexes. Traffic 1: 301–305.
27. Tortorella D, Gewurz BE, Furman MH, Schust DJ, Ploegh HL (2000) Viral
subversion of the immune system. Annu Rev Immunol 18: 861–926.
28. Wiertz EJ, Jones TR, Sun L, Bogyo M, Geuze HJ, et al. (1996) The human
cytomegalovirus US11 gene product dislocates MHC class I heavy chains from
the endoplasmic reticulum to the cytosol. Cell 84: 769–779.
29. Mueller B, Lilley BN, Ploegh HL (2006) SEL1L, the homologue of yeast Hrd3p,
is involved in protein dislocation from the mammalian ER. J Cell Biol 175:
261–270.
30. Mueller B, Klemm EJ, Spooner E, Claessen JH, Ploegh HL (2008) SEL1L
nucleates a protein complex required for dislocation of misfolded glycoproteins.
Proc Natl Acad Sci U S A 105: 12325–12330.
31. Baker BM, Tortorella D (2007) Dislocation of an endoplasmic reticulum
membrane glycoprotein involves the formation of partially dislocated ubiquitinated
polypeptides. J Biol Chem 282: 26845–26856.
32. Schulze A, Standera S, Buerger E, Kikkert M, van Voorden S, et al. (2005) The
ubiquitin-domain protein HERP forms a complex with components of the
endoplasmic reticulum associated degradation pathway. J Mol Biol 354:
1021–1027.
33. Burr ML, Cano F, Svobodova S, Boyle LH, Boname JM, et al. (2011) HRD1
and UBE2J1 target misfolded MHC class I heavy chains for endoplasmic
reticulum-associated degradation. Proc Natl Acad Sci U S A 108: 2034–2039.
34. Fiebiger E, Hirsch C, Vyas JM, Gordon E, Ploegh HL, et al. (2004) Dissection of
the dislocation pathway for type I membrane proteins with a new small molecule
inhibitor, eeyarestatin. Mol Biol Cell 15: 1635–1646.
35. Wang Q, Li L, Ye Y (2008) Inhibition of p97-dependent protein degradation by
Eeyarestatin I. J Biol Chem 283: 7445–7454.
36. Wang Q, Shinkre BA, Lee JG, Weniger MA, Liu Y, et al. (2010) The ERAD
inhibitor Eeyarestatin I is a bifunctional compound with a membrane-binding
domain and a p97/VCP inhibitory group. PLoS One 5: e15479.
37. Dolan BP, Li L, Veltri CA, Ireland CM, Bennink JR, et al. (2011) Distinct
pathways generate peptides from defective ribosomal products for CD8+ T cell
immunosurveillance. J Immunol 186: 2065–2072.
38. Cross BC, McKibbin C, Callan AC, Roboti P, Piacenti M, et al. (2009)
Eeyarestatin I inhibits Sec61-mediated protein translocation at the endoplasmic
reticulum. J Cell Sci 122: 4393–4400.
39. Ernst R, Claessen JHL, Mueller B, Sanyal S, Spooner E, et al. (2011) Enzymatic
blockade of the ubiquitin-proteasome pathway. PLoS Biol 8: e10000605.
40. Craiu A, Gaczynska M, Akopian T, Gramm CF, Fenteany G, et al. (1997)
Lactacystin and clasto-lactacystin beta-lactone modify multiple proteasome betasubunits
and inhibit intracellular protein degradation and major histocompatibility
complex class I antigen presentation. J Biol Chem 272: 13437–13445.
41. Tam PJ, Lingwood CA (2007) Membrane cytosolic translocation of verotoxin A1
subunit in target cells. Microbiology 153: 2700–2710.
42. Hudson TH, Neville DM, Jr. (1987) Temporal separation of protein toxin
translocation from processing events. J Biol Chem 262: 16484–16494.
43. Neville DM, Jr., Youle RJ (1982) Monoclonal antibody-ricin or ricin A chain
hybrids: kinetic analysis of cell killing for tumor therapy. Immunol Rev 62:
75–91.
44. Sandvig K, Ryd M, Garred O, Schweda E, Holm PK, et al. (1994) Retrograde
transport from the Golgi complex to the ER of both Shiga toxin and the
nontoxic Shiga B-fragment is regulated by butyric acid and cAMP. J Cell Biol
126: 53–64.
45. Sandvig K, Garred O, Prydz K, Kozlov JV, Hansen SH, et al. (1992)
Retrograde transport of endocytosed Shiga toxin to the endoplasmic reticulum.
Nature 358: 510–512.
46. Mallard F, Antony C, Tenza D, Salamero J, Goud B, et al. (1998) Direct
pathway from early/recycling endosomes to the Golgi apparatus revealed
through the study of shiga toxin B-fragment transport. J Cell Biol 143: 973–990.
47. Spooner RA, Watson P, Smith DC, Boal F, Amessou M, et al. (2008) The
secretion inhibitor Exo2 perturbs trafficking of Shiga toxin between endosomes
and the trans-Golgi network. Biochem J 414: 471–484.
48. Blewitt MG, Chung LA, London E (1985) Effect of pH on the conformation of
diphtheria toxin and its implications for membrane penetration. Biochemistry
24: 5458–5464.
49. Gallione CJ, Rose JK (1985) A single amino acid substitution in a hydrophobic
domain causes temperature-sensitive cell-surface transport of a mutant viral
glycoprotein. J Virol 54: 374–382.
50. Presley JF, Cole NB, Schroer TA, Hirschberg K, Zaal KJ, et al. (1997) ER-to-
Golgi transport visualized in living cells. Nature 389: 81–85.
51. Hammond C, Helenius A (1994) Quality control in the secretory pathway:
retention of a misfolded viral membrane glycoprotein involves cycling between
the ER, intermediate compartment, and Golgi apparatus. J Cell Biol 126: 41–52.
52. Donaldson JG, Lippincott-Schwartz J, Bloom GS, Kreis TE, Klausner RD
(1990) Dissociation of a 110-kD peripheral membrane protein from the Golgi
apparatus is an early event in brefeldin A action. J Cell Biol 111: 2295–2306.
53. Orci L, Tagaya M, Amherdt M, Perrelet A, Donaldson JG, et al. (1991)
Brefeldin A, a drug that blocks secretion, prevents the assembly of non-clathrincoated
buds on Golgi cisternae. Cell 64: 1183–1195.
54. Lippincott-Schwartz J, Yuan LC, Bonifacino JS, Klausner RD (1989) Rapid
redistribution of Golgi proteins into the ER in cells treated with brefeldin A:
evidence for membrane cycling from Golgi to ER. Cell 56: 801–813.
55. Chen SS, Huang AS (1986) Further characterization of the vesicular stomatitis
virus temperature-sensitive O45 mutant: intracellular conversion of the
glycoprotein to a soluble form. J Virol 59: 210–215.
56. Scheel J, Pepperkok R, Lowe M, Griffiths G, Kreis TE (1997) Dissociation of
coatomer from membranes is required for brefeldin A-induced transfer of Golgi
enzymes to the endoplasmic reticulum. J Cell Biol 137: 319–333.
57. Wang Q, Mora-Jensen H, Weniger MA, Perez-Galan P, Wolford C, et al. (2009)
ERAD inhibitors integrate ER stress with an epigenetic mechanism to activate
BH3-only protein NOXA in cancer cells. Proc Natl Acad Sci U S A 106:
2200–2205.
58. Kar R, Singha PK, Venkatachalam MA, Saikumar P (2009) A novel role for
MAP1 LC3 in nonautophagic cytoplasmic vacuolation death of cancer cells.
Oncogene 28: 2556–2568.
59. White J, Johannes L, Mallard F, Girod A, Grill S, et al. (1999) Rab6 coordinates
a novel Golgi to ER retrograde transport pathway in live cells. J Cell Biol 147:
743–760.
60. Monier S, Jollivet F, Janoueix-Lerosey I, Johannes L, Goud B (2002)
Characterization of novel Rab6-interacting proteins involved in endosome-to-
TGN transport. Traffic 3: 289–297.
61. Del Nery E, Miserey-Lenkei S, Falguieres T, Nizak C, Johannes L, et al. (2006)
Rab6A and Rab6A’ GTPases play non-overlapping roles in membrane
trafficking. Traffic 7: 394–407.
62. Chen A, AbuJarour RJ, Draper RK (2003) Evidence that the transport of ricin
to the cytoplasm is independent of both Rab6A and COPI. J Cell Sci 116:
3503–3510.
63. Amessou M, Fradagrada A, Falguieres T, Lord JM, Smith DC, et al. (2007)
Syntaxin 16 and syntaxin 5 are required for efficient retrograde transport of
several exogenous and endogenous cargo proteins. J Cell Sci 120: 1457–1468.
64. Sko¨nland SS, Walchli S, Utskarpen A, Wandinger-Ness A, Sandvig K (2007)
Phosphoinositide-regulated retrograde transport of ricin: crosstalk between
hVps34 and sorting nexins. Traffic 8: 297–309.
65. Dyve AB, Bergan J, Utskarpen A, Sandvig K (2009) Sorting nexin 8 regulates
endosome-to-Golgi transport. Biochem Biophys Res Commun 390: 109–114.
66. Bujny MV, Popoff V, Johannes L, Cullen PJ (2007) The retromer component
sorting nexin-1 is required for efficient retrograde transport of Shiga toxin from
early endosome to the trans Golgi network. J Cell Sci 120: 2010–2021.
67. Popoff V, Mardones GA, Bai SK, Chambon V, Tenza D, et al. (2009) Analysis
of articulation between clathrin and retromer in retrograde sorting on early
endosomes. Traffic 10: 1868–1880.
68. Smith DC, Spooner RA, Watson PD, Murray JL, Hodge TW, et al. (2006)
Internalized Pseudomonas exotoxin A can exploit multiple pathways to reach
the endoplasmic reticulum. Traffic 7: 379–393.
69. Chadfield MS, Hinton MH (2004) In vitro activity of nitrofuran derivatives on
growth and morphology of Salmonella enterica serotype Enteritidis. J Appl
Microbiol 96: 1002–1012.
70. Polycarpou-Schwarz M, Muller K, Denger S, Riddell A, Lewis J, et al. (2007)
Thanatop: a novel 5-nitrofuran that is a highly active, cell-permeable inhibitor of
topoisomerase II. Cancer Res 67: 4451–4458.
71. Maya JD, Bollo S, Nunez-Vergara LJ, Squella JA, Repetto Y, et al. (2003)
Trypanosoma cruzi: effect and mode of action of nitroimidazole and nitrofuran
derivatives. Biochem Pharmacol 65: 999–1006.
72. Letelier ME, Izquierdo P, Godoy L, Lepe AM, Faundez M (2004) Liver
microsomal biotransformation of nitro-aryl drugs: mechanism for potential
oxidative stress induction. J Appl Toxicol 24: 519–525.
73. Lakkaraju AK, Luyet PP, Parone P, Falguieres T, Strub K (2007) Inefficient
targeting to the endoplasmic reticulum by the signal recognition particle elicits
selective defects in post-ER membrane trafficking. Exp Cell Res 313: 834–847.
74. Clague MJ, Urbe S (2006) Endocytosis: the DUB version. Trends Cell Biol 16:
551–559.
75. Wakana Y, Takai S, Nakajima K, Tani K, Yamamoto A, et al. (2008) Bap31 is
an itinerant protein that moves between the peripheral endoplasmic reticulum
(ER) and a juxtanuclear compartment related to ER-associated Degradation.
Mol Biol Cell 19: 1825–1836.
76. Hosokawa N, You Z, Tremblay LO, Nagata K, Herscovics A (2007) Stimulation
of ERAD of misfolded null Hong Kong alpha1-antitrypsin by Golgi alpha1,2-
mannosidases. Biochem Biophys Res Commun 362: 626–632.
77. Vashist S, Kim W, Belden WJ, Spear ED, Barlowe C, et al. (2001) Distinct
retrieval and retention mechanisms are required for the quality control of
endoplasmic reticulum protein folding. J Cell Biol 155: 355–368.
78. Fu L, Sztul E (2003) Traffic-independent function of the Sar1p/COPII
machinery in proteasomal sorting of the cystic fibrosis transmembrane
conductance regulator. J Cell Biol 160: 157–163.
79. Aghi M, Kramm CM, Chou TC, Breakefield XO, Chiocca EA (1998)
Synergistic anticancer effects of ganciclovir/thymidine kinase and 5-fluorocytosine/
cytosine deaminase gene therapies. J Natl Cancer Inst 90: 370–380.
80. Chiu CF, Ghanekar Y, Frost L, Diao A, Morrison D, et al. (2008) ZFPL1, a
novel ring finger protein required for cis-Golgi integrity and efficient ER-to-
Golgi transport. Embo J 27: 934–947.
81. Schagger H, von Jagow G (1987) Tricine-sodium dodecyl sulfate-polyacrylamide
gel electrophoresis for the separation of proteins in the range from 1 to 100 kDa.
Anal Biochem 166: 368–379.