The peroxisomal protein import machinery displays a preference for monomeric substrates

Open Biology

Volumn 5, Issue 4, 2015, Pages

  Freitas, Marta O ^a   Francisco, Tânia ^a   Rodrigues, Tony A ^a,b   Lismont, Celien ^d   Domingues, Pedro ^c   Pinto, Manuel P ^a   Grou, Cláudia P ^a   Fransen, Marc ^d   Azevedo, Jorge E ^a,b  

^a UNIVERSITY OF PORTO   (Portugal)

^b UNIVERSITY OF PORTO   (Portugal)

^c UNIVERSITY OF AVEIRO   (Portugal)

^d UNIVERSITY OF LEUVEN   (Belgium)

Author keywords

Acyl CoA oxidase; Docking translocation machinery; Peroxisomes; PEX5; Protein import; Urate oxidase

Indexed keywords

ACYL COENZYME A OXIDASE; CELL RECEPTOR; PEROXISOME-TARGETING SIGNAL 1 RECEPTOR; PROTEIN BINDING; URATE OXIDASE;

ANIMAL; BIOLOGICAL MODEL; CHEMISTRY; CHLOROCEBUS AETHIOPS; COS CELL LINE; CYTOSOL; FLUORESCENCE MICROSCOPY; GENETICS; HUMAN; METABOLISM; MOUSE; MUTATION; PEROXISOME; PROTEIN MULTIMERIZATION; PROTEIN TRANSPORT; RAT; SIGNAL TRANSDUCTION; WESTERN BLOTTING;

ACYL-COA OXIDASE; ANIMALS; BLOTTING, WESTERN; CERCOPITHECUS AETHIOPS; COS CELLS; CYTOSOL; HUMANS; MICE; MICROSCOPY, FLUORESCENCE; MODELS, BIOLOGICAL; MUTATION; PEROXISOMES; PROTEIN BINDING; PROTEIN MULTIMERIZATION; PROTEIN TRANSPORT; RATS; RECEPTORS, CYTOPLASMIC AND NUCLEAR; SIGNAL TRANSDUCTION; URATE OXIDASE;

EID: 84934927612     PISSN: None     EISSN: 20462441     Source Type: Journal    
DOI: 10.1098/rsob.140236     Document Type: Article

References (85)

1. Wanders RJA. 2014 Metabolic functions of peroxisomes in health and disease. Biochimie 98, 36-44. (doi:10.1016/j.biochi.2013.08.022)
2. Van Veldhoven PP. 2010 Biochemistry and genetics of inherited disorders of peroxisomal fatty acid metabolism. J. Lipid Res. 51, 2863-2895. (doi:10. 1194/jlr. R005959)
3. Hu J, Baker A, Bartel B, Linka N, Mullen RT, Reumann S, Zolman BK. 2012 Plant peroxisomes: biogenesis and function. Plant Cell 24, 2279-2303. (doi:10.1105/tpc.112.096586)
4. Michels PAM, Bringaud F, Herman M, Hannaert V. 2006 Metabolic functions of glycosomes in trypanosomatids. Biochim. Biophys. Acta 1763, 1463-1477. (doi:10.1016/j.bbamcr.2006.08.019)
5. Wiese S et al. 2007 Proteomics characterization of mouse kidney peroxisomes by tandem mass spectrometry and protein correlation profiling. Mol. Cell. Proteomics 6, 2045-2057. (doi:10.1074/mcp. M700169-MCP200)
6. Kikuchi M, Hatano N, Yokota S, Shimozawa N, Imanaka T, Taniguchi H. 2004 Proteomic analysis of rat liver peroxisome: presence of peroxisomespecific isozyme of Lon protease. J. Biol. Chem. 279, 421-428. (doi:10.1074/jbc. M305623200)
7. Lazarow PB, Fujiki Y. 1985 Biogenesis of peroxisomes. Annu. Rev. Cell Biol. 1, 489-530. (doi:10.1146/annurev. cb.01.110185. 002421)
8. Brocard C, Hartig A. 2006 Peroxisome targeting signal 1: is it really a simple tripeptide? Biochim. Biophys. Acta 1763, 1565-1573. (doi:10.1016/j. bbamcr.2006.08.022)
9. Gould SJ, Keller GA, Hosken N, Wilkinson J, Subramani S. 1989 A conserved tripeptide sorts proteins to peroxisomes. J. Cell Biol. 108, 1657-1664. (doi:10.1083/jcb.108.5.1657)
10. Kunze M, Neuberger G, Maurer-Stroh S, Ma J, Eck T, Braverman N, Schmid JA, Eisenhaber F, Berger J. 2011 Structural requirements for interaction of peroxisomal targeting signal 2 and its receptor PEX7. J. Biol. Chem. 286, 45 048-45 062. (doi:10.1074/jbc. M111.301853)
11. Swinkels BW, Gould SJ, Bodnar AG, Rachubinski RA, Subramani S. 1991 A novel, cleavable peroxisomal targeting signal at the amino-terminus of the rat 3-ketoacyl-CoA thiolase. EMBO J. 10, 3255-3262.
12. Lazarow PB. 2006 The import receptor Pex7p and the PTS2 targeting sequence. Biochim. Biophys. Acta 1763, 1599-1604. (doi:10.1016/j.bbamcr. 2006.08.011)
13. Francisco T, Rodrigues TA, Pinto MP, Carvalho AF, Azevedo JE, Grou CP. 2014 Ubiquitin in the peroxisomal protein import pathway. Biochimie 98, 29-35. (doi:10.1016/j.biochi.2013.08.003)
14. Baker A, Paudyal R. 2014 The life of the peroxisome: from birth to death. Curr. Opin. Plant Biol. 22, 39-47. (doi:10.1016/j.pbi.2014.09.003)
15. Platta HW, Hagen S, Reidick C, Erdmann R. 2014 The peroxisomal receptor dislocation pathway: to the exportomer and beyond. Biochimie 98, 16-28. (doi:10.1016/j.biochi.2013.12.009)
16. Liu X, Ma C, Subramani S. 2012 Recent advances in peroxisomal matrix protein import. Curr. Opin. Cell Biol. 24, 484-489. (doi:10.1016/j.ceb.2012.05.003)
17. Dodt G, Gould SJ. 1996 Multiple PEX genes are required for proper subcellular distribution and stability of Pex5p, the PTS1 receptor: evidence that PTS1 protein import is mediated by a cycling receptor. J. Cell Biol. 135, 1763-1774. (doi:10. 1083/jcb.135.6.1763)
18. Gatto G, Geisbrecht B. 2000 Peroxisomal targeting signal-1 recognition by the TPR domains of human PEX5. Nat. Struct. Biol. 7, 1091-1095. (doi:10.1038/81930)
19. Braverman N, Steel G, Obie C, Moser A, Moser H, Gould SJ, Valle D. 1997 Human PEX7 encodes the peroxisomal PTS2 receptor and is responsible for rhizomelic chondrodysplasia punctata. Nat. Genet. 15, 369-376. (doi:10.1038/ng0497-369)
20. Otera H et al. 1998 Peroxisome targeting signal type 1 (PTS1) receptor is involved in import of both PTS1 and PTS2: studies with PEX5-defective CHO cell mutants. Mol. Cell. Biol. 18, 388-399.
21. Woodward AW, Bartel B. 2005 The Arabidopsis peroxisomal targeting signal type 2 receptor PEX7 is necessary for peroxisome function and dependent on PEX5. Mol. Biol. Cell 16, 573-583. (doi:10.1091/mbc. E04-05-0422)
22. Gouveia AM, Guimaraes CP, Oliveira ME, Reguenga C, Sa-Miranda C, Azevedo JE. 2003 Characterization of the peroxisomal cycling receptor, Pex5p, using a cell-free in vitro import system. J. Biol. Chem. 278, 226-232. (doi:10.1074/jbc. M209498200)
23. Francisco T, Rodrigues TA, Freitas MO, Grou CP, Carvalho AF, Sá-Miranda C, Pinto MP, Azevedo JE. 2013 A cargo-centered perspective on the PEX5-mediated peroxisomal protein import pathway. J. Biol. Chem. 288, 29 151-29 159. (doi:10.1074/jbc. M113.487140)
24. Rodrigues TA, Alencastre IS, Francisco T, Brites P, Fransen M, Grou CP, Azevedo JE. 2014 A PEX7-centered perspective on the peroxisomal targeting signal type 2-mediated protein import pathway. Mol. Cell. Biol. 34, 2917-2928. (doi:10.1128/MCB. 01727-13)
25. Reguenga C, Oliveira ME, Gouveia AM, Sá-Miranda C, Azevedo JE. 2001 Characterization of the mammalian peroxisomal import machinery: Pex2p, Pex5p, Pex12p, and Pex14p are subunits of the same protein assembly. J. Biol. Chem. 276, 29 935-29 942. (doi:10.1074/jbc. M104114200)
26. Oeljeklaus S et al. 2012 Identification of core components and transient interactors of the peroxisomal importomer by dual-track stable isotope labeling with amino acids in cell culture analysis. J. Proteome Res. 11, 2567-2580. (doi:10. 1021/pr3000333)
27. Gouveia AM, Guimarães CP, Oliveira ME, Sá-Miranda C, Azevedo JE. 2003 Insertion of Pex5p into the peroxisomal membrane is cargo protein-dependent. J. Biol. Chem. 278, 4389-4392. (doi:10.1074/jbc. C200650200)
28. Alencastre IS, Rodrigues TA, Grou CP, Fransen M, Sá-Miranda C, Azevedo JE. 2009 Mapping the cargo protein membrane translocation step into the PEX5 cycling pathway. J. Biol. Chem. 284, 27 243-27 251. (doi:10.1074/jbc. M109.032565)
29. Oliveira ME, Gouveia AM, Pinto RA, Sá-Miranda C, Azevedo JE. 2003 The energetics of Pex5p-mediated peroxisomal protein import. J. Biol. Chem. 278, 39 483-39 489. (doi:10.1074/jbc. M305089200)
30. Azevedo JE, Costa-Rodrigues J, Guimarães CP, Oliveira ME, Sá-Miranda C. 2004 Protein translocation across the peroxisomal membrane. Cell Biochem. Biophys. 41, 451-468. (doi:10.1385/CBB:41:3:451)
31. Williams C, van den Berg M, Sprenger RR, Distel B. 2007 A conserved cysteine is essential for Pex4pdependent ubiquitination of the peroxisomal import receptor Pex5p. J. Biol. Chem. 282, 22 534-22 543. (doi:10.1074/jbc. M702038200)
32. Carvalho AF, Pinto MP, Grou CP, Alencastre IS, Fransen M, Sá-Miranda C, Azevedo JE. 2007 Ubiquitination of mammalian Pex5p, the peroxisomal import receptor. J. Biol. Chem. 282, 31 267-31 272. (doi:10.1074/jbc. M706325200)
33. Miyata N, Fujiki Y. 2005 Shuttling mechanism of peroxisome targeting signal type 1 receptor Pex5?: ATP-independent import and ATP-dependent export. Mol. Cell. Biol. 25, 10 822-10 832. (doi:10. 1128/MCB.25.24.10822)
34. Platta HW, Grunau S, Rosenkranz K, Girzalsky W, Erdmann R. 2005 Functional role of the AAA peroxins in dislocation of the cycling PTS1 receptor back to the cytosol. Nat. Cell Biol. 7, 817-822. (doi:10.1038/ncb1281)
35. Debelyy MO, Platta HW, Saffian D, Hensel A, Thoms S, Meyer HE, Warscheid B, Girzalsky W, Erdmann R. 2011 Ubp15p, a ubiquitin hydrolase associated with the peroxisomal export machinery. J. Biol. Chem. 286, 28 223-28 234. (doi:10.1074/jbc. M111.238600)
36. Grou CP, Carvalho AF, Pinto MP, Huybrechts SJ, Sá-Miranda C, Fransen M, Azevedo JE. 2009 Properties of the ubiquitin-pex5p thiol ester conjugate. J. Biol. Chem. 284, 10 504-10 513. (doi:10.1074/jbc. M808978200)
37. Grou CP et al. 2012 Identification of ubiquitinspecific protease 9X (USP9X) as a deubiquitinase acting on the ubiquitin-peroxin 5 (PEX5) thioester conjugate. J. Biol. Chem. 287, 12 815-12 827. (doi:10.1074/jbc. M112.340158)
38. Otera H, Fujiki Y. 2012 Pex5p imports folded tetrameric catalase by interaction with Pex13p. Traffic 13, 1364-1377. (doi:10.1111/j.1600-0854. 2012.01391.x)
39. Glover JR, Andrews DW, Rachubinski RA. 1994 Saccharomyces cerevisiae peroxisomal thiolase is imported as a dimer. Proc. Natl Acad. Sci. USA 91, 10 541-10 545. (doi:10.1073/pnas.91.22.10541)
40. McNew JA, Goodman JM. 1994 An oligomeric protein is imported into peroxisomes in vivo. J. Cell Biol. 127, 1245-1257. (doi:10.1083/jcb.127.5.1245)
41. Yang X, Purdue PE, Lazarow PB. 2001 Eci1p uses a PTS1 to enter peroxisomes: either its own or that of a partner, Dci1p. Eur. J. Cell Biol. 80, 126-138. (doi:10.1078/0171-9335-00144)
42. Schueren F, Lingner T, George R, Hofhuis J, Dickel C, Gärtner J, Thoms S. 2014 Peroxisomal lactate dehydrogenase is generated by translational read through in mammals. eLife 3, e3640. (doi:10.7554/eLife.03640)
43. Lee MS, Mullen RT, Trelease RN. 1997 Oilseed isocitrate lyases lacking their essential type 1 peroxisomal targeting signal are piggybacked to glyoxysomes. Plant Cell 9, 185-197. (doi:10.1105/tpc.9.2.185)
44. Islinger M, Li KW, Seitz J, Völkl A, Lüers GH. 2009 Hitchhiking of Cu/Zn superoxide dismutase to peroxisomes-evidence for a natural piggyback import mechanism in mammals. Traffic 10, 1711-1721. (doi:10.1111/j.1600-0854.2009.00966.x)
45. Stewart MQ, Esposito RD, Gowani J, Goodman JM. 2001 Alcohol oxidase and dihydroxyacetone synthase, the abundant peroxisomal proteins of methylotrophic yeasts, assemble in different cellular compartments. J. Cell Sci. 114, 2863-2868.
46. Titorenko VI, Nicaud J-M, Wang H, Chan H, Rachubinski RA. 2002 Acyl-CoA oxidase is imported as a heteropentameric, cofactor-containing complex into peroxisomes of Yarrowia lipolytica. J. Cell Biol. 156, 481-494. (doi:10.1083/jcb.200111075)
47. Gunkel K, Veenhuis M, van der Klei IJ. 2005 Protein translocation machineries: how organelles bring in matrix proteins. FEMS Yeast Res. 5, 1037-1045. (doi:10.1016/j.femsyr.2005.03.004)
48. Gould SJ, Collins CS. 2002 Opinion: peroxisomalprotein import: is it really that complex? Nat. Rev. Mol. Cell Biol. 3, 382-389. (doi:10.1038/nrm807)
49. Smith JJ, Aitchison JD. 2013 Peroxisomes take shape. Nat. Rev. Mol. Cell Biol. 14, 803-817. (doi:10.1038/nrm3700)
50. Kumar S, Kawałek A, van der Klei IJ. 2014 Peroxisomal quality control mechanisms. Curr. Opin. Microbiol. 22 C, 30-37. (doi:10.1016/j.mib. 2014.09.009)
51. Lodish HF. 2013 Molecular cell biology. New York, NY: W. H. Freeman.
52. Lazarow PB, de Duve C. 1973 The synthesis and turnover of rat liver peroxisomes. V. Intracellular pathway of catalase synthesis. J. Cell Biol. 59, 507-524. (doi:10.1083/jcb.59.2.507)
53. Goodman JM, Scott CW, Donahue PN, Atherton JP. 1984 Alcohol oxidase assembles post-translationally into the peroxisome of Candida boidinii. J. Biol. Chem. 259, 8485-8493.
54. Crookes WJ, Olsen LJ. 1998 The effects of chaperones and the influence of protein assembly on peroxisomal protein import. J. Biol. Chem. 273, 17 236-17 242. (doi:10.1074/jbc.273.27.17236)
55. Brocard C, Jedeszko C, Song HC, Terlecky SR, Walton PA. 2003 Protein structure and import into the peroxisomal matrix. Traffic 4, 74-82. (doi:10.1034/j.1600-0854.2003.40203.x)
56. Tanaka N, Aoki K-I, Ishikura S, Nagano M, Imamura Y, Hara A, Nakamura KT. 2008 Molecular basis for peroxisomal localization of tetrameric carbonyl reductase. Structure 16, 388-397. (doi:10.1016/j. str.2007.12.022)
57. Luo B, Norris C, Bolstad ESD, Knecht DA, Grant DF. 2008 Protein quaternary structure and expression levels contribute to peroxisomal-targetingsequence-1-mediated peroxisomal import of human soluble epoxide hydrolase. J. Mol. Biol. 380, 31-41. (doi:10.1016/j.jmb.2008.04.064)
58. Faber KN, van Dijk R, Keizer-Gunnink I, Koek A, van der Klei IJ, Veenhuis M. 2002 Import of assembled PTS1 proteins into peroxisomes of the yeast Hansenula polymorpha: yes and no! Biochim. Biophys. Acta 1591, 157-162. (doi:10.1016/S0167-4889 (02) 00274-4)
59. Grou CP, Carvalho AF, Pinto MP, Alencastre IS, Rodrigues TA, Freitas MO, Francisco T, Sá-Miranda C, Azevedo JE. 2009 The peroxisomal protein import machinery-a case report of transient ubiquitination with a new flavor. Cell. Mol. Life Sci. 66, 254-262. (doi:10.1007/s00018-008-8415-5)
60. Costa-Rodrigues J, Carvalho AF, Fransen M, Hambruch E, Schliebs W, Sá-Miranda C, Azevedo JE. 2005 Pex5p, the peroxisomal cycling receptor, is a monomeric nonglobular protein. J. Biol. Chem. 280, 24 404-24 411. (doi:10.1074/jbc. M501985200)
61. Freitas MO et al. 2011 PEX5 binds monomeric catalase blocking its tetramerization, and releases it upon binding the N-terminal domain of PEX14. J. Biol. Chem. 286, 40 509-40 519. (doi:10.1074/jbc. M111.287201)
62. Hashimoto T. 1999 Peroxisomal beta-oxidation enzymes. Neurochem. Res. 24, 551-563. (doi:10. 1023/A:1022540030918)
63. Kratzer JT, Lanaspa MA, Murphy MN, Cicerchi C, Graves CL, Tipton PA, Ortlund EA, Johnson RJ, Gaucher EA. 2014 Evolutionary history and metabolic insights of ancient mammalian uricases. Proc. Natl Acad. Sci. USA 111, 3763-3768. (doi:10. 1073/pnas.1320393111)
64. Leighton F, Poole B, Lazarow PB, De Duve C. 1969 The synthesis and turnover of rat liver peroxisomes. I. Fractionation of peroxisome proteins. J. Cell Biol. 41, 521-535. (doi:10.1083/jcb.41.2.521)
65. Miura S, Mori M, Takiguchi M, Tatibana M, Furuta S, Miyazawa S, Hashimoto T. 1984 Biosynthesis and intracellular transport of enzymes of peroxisomal beta-oxidation. J. Biol. Chem. 259, 6397-6402.
66. Miyazawa S, Hayashi H, Hijikata M, Ishii N, Furuta S, Kagamiyama H, Osumi T, Hashimoto T. 1987 Complete nucleotide sequence of cDNA and predicted amino acid sequence of rat acyl-CoA oxidase. J. Biol. Chem. 262, 8131-8137.
67. Schram AW, Strijland A, Hashimoto T, Wanders RJ, Schutgens RB, van den Bosch H, Tager JM. 1986 Biosynthesis and maturation of peroxisomal betaoxidation enzymes in fibroblasts in relation to the Zellweger syndrome and infantile Refsum disease. Proc. Natl Acad. Sci. USA 83, 6156-6158. (doi:10. 1073/pnas.83.16.6156)
68. Szilard RK, Rachubinski RA. 2000 Tetratricopeptide repeat domain of Yarrowia lipolytica Pex5p is essential for recognition of the type 1 peroxisomal targeting signal but does not confer full biological activity on Pex5p. Biochem. J. 346, 177-184. (doi:10.1042/0264-6021:3460177)
69. Dodt G, Braverman N, Wong C, Moser A, Moser HW, Watkins P, Valle D, Gould SJ. 1995 Mutations in the PTS1 receptor gene, PXR1, define complementation group 2 of the peroxisome biogenesis disorders. Nat. Genet. 9, 115-125. (doi:10.1038/ng0295-115)
70. Hageman J, Vos MJ, van Waarde MAW H, Kampinga HH. 2007 Comparison of intra-organellar chaperone capacity for dealing with stress-induced protein unfolding. J. Biol. Chem. 282, 34 334-34 345. (doi:10.1074/jbc. M703876200)
71. Bartoszewska M, Williams C, Kikhney A, Opaliński Ł, van Roermund CWT, de Boer R, Veenhuis M, van der Klei IJ. 2012 Peroxisomal proteostasis involves a Lon family protein that functions as protease and chaperone. J. Biol. Chem. 287, 27 380-27 395. (doi:10.1074/jbc. M112.381566)
72. Léon S, Goodman JM, Subramani S. 2006 Uniqueness of the mechanism of protein import into the peroxisome matrix: transport of folded, cofactorbound and oligomeric proteins by shuttling receptors. Biochim. Biophys. Acta 1763, 1552-1564. (doi:10.1016/j.bbamcr.2006.08.037)
73. Danpure CJ. 2006 Primary hyperoxaluria type 1: AGT mistargeting highlights the fundamental differences between the peroxisomal and mitochondrial protein import pathways. Biochim. Biophys. Acta 1763, 1776-1784. (doi:10.1016/j.bbamcr.2006.08.021)
74. Palosaari PM, Hiltunen JK. 1990 Peroxisomal bifunctional protein from rat liver is a trifunctional enzyme possessing 2-enoyl-CoA hydratase, 3-hydroxyacyl-CoA dehydrogenase, and delta 3, delta 2-enoyl-CoA isomerase activities. J. Biol. Chem. 265, 2446-2449.
75. Klootwijk ED et al. 2014 Mistargeting of peroxisomal EHHADH and inherited renal Fanconi's syndrome. N. Engl. J. Med. 370, 129-138. (doi:10. 1056/NEJMoa1307581)
76. Pey AL, Salido E, Sanchez-Ruiz JM. 2011 Role of low native state kinetic stability and interaction of partially unfolded states with molecular chaperones in the mitochondrial protein mistargeting associated with primary hyperoxaluria. Amino Acids 41, 1233-1245. (doi:10.1007/s00726-010-0801-2)
77. Chacinska A, Koehler CM, Milenkovic D, Lithgow T, Pfanner N. 2009 Importing mitochondrial proteins: machineries and mechanisms. Cell 138, 628-644. (doi:10.1016/j.cell.2009.08.005)
78. Carvalho AF, Costa-Rodrigues J, Correia I, Costa Pessoa J, Faria TQ, Martins CL, Fransen M, Sá-Miranda C, Azevedo JE. 2006 The N-terminal half of the peroxisomal cycling receptor Pex5p is a natively unfolded domain. J. Mol. Biol. 356, 864-875. (doi:10.1016/j.jmb.2005.12.002)
79. Carvalho AF, Grou CP, Pinto MP, Alencastre IS, Costa-Rodrigues J, Fransen M, Sá-Miranda C, Azevedo JE. 2007 Functional characterization of two missense mutations in Pex5p - C11S and N526K. Biochim. Biophys. Acta 1773, 1141-1148. (doi:10.1016/j. bbamcr.2007.04.011)
80. Huybrechts SJ, Van Veldhoven PP, Brees C, Mannaerts GP, Los GV, Fransen M. 2009 Peroxisome dynamics in cultured mammalian cells. Traffic 10, 1722-1733. (doi:10.1111/j.1600-0854.2009. 00970.x)
81. Amery L, Fransen M, De Nys K, Mannaerts GP, Van Veldhoven PP. 2000 Mitochondrial and peroxisomal targeting of 2-methylacyl-CoA racemase in humans. J. Lipid Res. 41, 1752-1759.
82. Nordgren M, Wang B, Apanasets O, Brees C, Veldhoven PP, Fransen M. 2012 Potential limitations in the use of KillerRed for fluorescence microscopy. J. Microsc. 245, 229-235. (doi:10.1111/j.1365-2818.2011.03564.x)
83. Hartl F-U, Just WW, Koster A, Schimassek H. 1985 Improved isolation and purification of rat liver peroxisomes by combined rate zonal and equilibrium density centrifugation. Arch. Biochem. Biophys. 237, 124-134. (doi:10.1016/0003-9861 (85) 90261-9)
84. Gouveia AM, Reguenga C, Oliveira ME, Eckerskorn C, Sá-Miranda C, Azevedo JE. 1999 Alkaline density gradient floatation of membranes: polypeptide composition of the mammalian peroxisomal membrane. Anal. Biochem. 274, 270-277. (doi:10. 1006/abio.1999.4281)
85. Baumgart E, Vanhooren JC, Fransen M, Van Leuven F, Fahimi HD, Van Veldhoven PP, Mannaerts GP. 1996 Molecular cloning and further characterization of rat peroxisomal trihydroxycoprostanoyl-CoA oxidase. Biochem. J. 320, 115-121.