A Structural View of Xenophagy, a Battle between Host and Microbes

다국어 초록 (Multilingual Abstract)

The cytoplasm in mammalian cells is a battlefield between the host and invading microbes. Both the living organisms have evolved unique strategies for their survival. The host utilizes a specialized autophagy system, xenophagy, for the clearance of invading pathogens, whereas bacteria secrete proteins to defend and escape from the host xenophagy. Several molecules have been identified and their structural investigation has enabled the comprehension of these mechanisms at the molecular level. In this review, we focus on one example of host autophagy and the other of bacterial defense: the autophagy receptor, NDP52, in conjunction with the sugar receptor, galectin-8, plays a critical role in targeting the autophagy machinery against Sal-monella; and the cysteine protease, RavZ secret-ed by Legionella pneumophila cleaves the LC3-PE on the phagophore membrane. The structure-function relationships of these two examples and the directions of future research will be dis-cussed.

참고문헌 (Reference)

1 Chen, W., "Unusual regioversatility of acetyltransferase Eis, a cause of drug resistance in XDR-TB" 108 : 9804-9808, 2011

2 Manzanillo, P.S., "The ubiquitin ligase parkin mediates resistance to intracellular pathogens" 501 : 512-516, 2013

3 Zhang, R.G., "The three-dimensional crystal structure of cholera toxin" 251 : 563-573, 1995

4 Satoo, K., "The structure of Atg4B-LC3 complex reveals the mechanism of LC3 processing and delipidation during autophagy" 28 : 1341-1350, 2009

5 Boyle, K.B., "The role of 'eat-me' signals and autophagy cargo receptors in innate immunity" 16 : 339-348, 2013

6 Thurston, T.L., "The TBK1 adaptor and autophagy receptor NDP52 restricts the proliferation of ubiquitin-coated bacteria" 10 : 1215-1221, 2009

7 Choy, A., "The Legionella effector RavZ inhibits host autophagy through irreversible Atg8 deconjugation" 338 : 1072-1076, 2012

8 Horenkamp, F.A., "The Legionella anti-autophagy effector RavZ targets the autophagosome via PI3P- and curvature-sensing motifs" 34 : 569-576, 2015

9 Huett, A., "The LRR and RING domain protein LRSAM1 is an E3 ligase crucial for ubiquitin-dependent autophagy of intracellular Salmonella Typhimurium" 12 : 778-790, 2012

10 Kwon, D.H., "The 1:2 complex between RavZ and LC3 reveals a mechanism for deconjugation of LC3 on the phagophore membrane" 13 : 70-81, 2017

11 김준회, "Swapping of interaction partners with ATG5 for autophagosome maturation" 생화학분자생물학회 48 (48): 129-130, 2015

12 Hong, S.B., "Structure of the autophagic E2 enzyme Atg10" 68 : 1409-1417, 2012

13 Neves, D., "Structure of internalin InlK from the human pathogen Listeria monocytogenes" 425 : 4520-4529, 2013

14 Shen, Y., "Structure of anthrax edema factor-calmodulinadenosine 5'-(alpha,beta-methylene)-triphosphate complex reveals an alternative mode of ATP binding to the catalytic site" 317 : 309-314, 2004

15 Kim, K.H., "Structure of Mycobacterium smegmatis Eis in complex with paromomycin" 70 : 1173-1179, 2014

16 김병원, "Structure biology of selective autophagy receptors" 생화학분자생물학회 49 (49): 73-80, 2016

17 Renshaw, P.S., "Structure and function of the complex formed by the tuberculosis virulence factors CFP-10 and ESAT-6" 24 : 2491-2498, 2005

18 Fan, E., "Structural biology and structure-based inhibitor design of cholera toxin and heat-labile enterotoxin" 294 : 217-223, 2004

19 Kim, B.W., "Structural basis for recognition of autophagic receptor NDP52 by the sugar receptor galectin-8" 4 : 1613-, 2013

20 Germane, K.L., "Structural and functional studies indicate that Shigella VirA is not a protease and does not directly destabilize microtubules" 47 : 10241-10243, 2008

21 Li, S., "Sterical hindrance promotes selectivity of the autophagy cargo receptor NDP52 for the danger receptor galectin-8 in antibacterial autophagy" 6 : ra9-, 2013

22 Rahighi, S., "Selectivity of the ubiquitin-binding modules" 586 : 2705-2710, 2012

23 Svenning, S., "Selective autophagy" 55 : 79-92, 2013

24 Perrin, A.J., "Recognition of bacteria in the cytosol of Mammalian cells by the ubiquitin system" 14 : 806-811, 2004

25 Liu, L., "Receptormediated mitophagy in yeast and mammalian systems" 24 : 787-795, 2014

26 Behrends, C., "Receptor proteins in selective autophagy" 2012 : 673290-, 2012

27 He, H., "Post-translational modifications of three members of the human MAP1LC3 family and detection of a novel type of modification for MAP1LC3B" 278 : 29278-29287, 2003

28 Davis, J., "Novel fold of VirA, a type III secretion system effector protein from Shigella flexneri" 17 : 2167-2173, 2008

29 Holmner, A., "Novel binding site identified in a hybrid between cholera toxin and heat-labile enterotoxin: 1.9 $\AA$ crystal structure reveals the details" 12 : 1655-1667, 2004

30 Shin, D.M., "Mycobacterium tuberculosis eis regulates autophagy, inflammation, and cell death through redox-dependent signaling" 6 : e1001230-, 2010

31 Kim, K.H., "Mycobacterium tuberculosis Eis protein initiates suppression of host immune responses by acetylation of DUSP16/MKP-7" 109 : 7729-7734, 2012

32 Yoshii, S.R., "Monitoring and Measuring Autophagy" 18 : 1865-, 2017

33 Farre, J.C., "Mechanistic insights into selective autophagy pathways: lessons from yeast" 17 : 537-552, 2016

34 Zaffagnini, G., "Mechanisms of selective autophagy" 428 : 1714-1724, 2016

35 Pantoom, S., "Lift and cut: Anti-host autophagy mechanism of Legionella pneumophila" 13 : 1467-1469, 2017

36 Ng, A., "Leucine-rich repeat (LRR) proteins: integrators of pattern recognition and signaling in immunity" 7 : 1082-1084, 2011

37 Noad, J., "LUBAC-synthesized linear ubiquitin chains restrict cytosol-invading bacteria by activating autophagy and NF-kappaB" 2 : 17063-, 2017

38 Celli, J., "LRSAM1, an E3 Ubiquitin ligase with a sense for bacteria" 12 : 735-736, 2012

39 Heckmann, B.L., "LC3-Associated Phagocytosis and Inflammation" 429 : 3561-3576, 2017

40 Hong, S.B., "Insights into noncanonical E1 enzyme activation from the structure of autophagic E1 Atg7 with Atg8" 18 : 1323-1330, 2011

41 Kim, J.H., "Insights into autophagosome maturation revealed by the structures of ATG5 with its interacting partners" 11 : 75-87, 2015

42 Ng, A.C., "Human leucine-rich repeat proteins: a genome-wide bioinformatic categorization and functional analysis in innate immunity" 108 (108): 4631-4638, 2011

43 Klionsky, D.J., "Guidelines for the use and interpretation of assays for monitoring autophagy (3rd edition)" 12 : 1-222, 2016

44 Thurston, T.L., "Galectin 8 targets damaged vesicles for autophagy to defend cells against bacterial invasion" 482 : 414-418, 2012

45 Ogawa, M., "Escape of intracellular Shigella from autophagy" 307 : 727-731, 2005

46 Sorbara, M.T., "Emerging themes in bacterial autophagy" 23 : 163-170, 2015

47 Yang, A., "Elucidation of the antiautophagy mechanism of the Legionella effector RavZ using semisynthetic LC3 proteins" 6 : e23905-, 2017

48 Levine, B, "Eating oneself and uninvited guests: autophagyrelated pathways in cellular defense" 120 : 159-162, 2005

49 Nakatogawa, H., "Dynamics and diversity in autophagy mechanisms: lessons from yeast" 10 : 458-467, 2009

50 Klionsky, D.J., "Dynamic regulation of macroautophagy by distinctive ubiquitin-like proteins" 21 : 336-345, 2014

51 Gangwer, K.A., "Crystal structure of the Helicobacter pylori vacuolating toxin p55 domain" 104 : 16293-16298, 2007

52 Merritt, E.A., "Crystal structure of cholera toxin B-pentamer bound to receptor GM1 pentasaccharide" 3 : 166-175, 1994

53 Santelli, E., "Crystal structure of a complex between anthrax toxin and its host cell receptor" 430 : 905-908, 2004

54 지창훈, "Crosstalk and Interplay between the Ubiquitin-Proteasome System and Autophagy" 한국분자세포생물학회 40 (40): 441-449, 2017

55 Tattoli, I., "Bacterial autophagy: the trigger, the target and the timing" 8 : 1848-1850, 2012

56 Huang, J., "Bacteria-autophagy interplay: a battle for survival" 12 : 101-114, 2014

57 Maruyama, T., "Autophagy-regulating protease Atg4: structure, function, regulation and inhibition" 71 : 72-78, 2018

58 Levine, B., "Autophagy wins the 2016 Nobel Prize in Physiology or Medicine: Breakthroughs in baker's yeast fuel advances in biomedical research" 114 : 201-205, 2017

59 Mizushima, N, "Autophagy in protein and organelle turnover" 76 : 397-402, 2011

60 Levine, B., "Autophagy in immunity and inflammation" 469 : 323-335, 2011

61 나지훈, "Autophagy in Neurodegenerative Diseases: From Mechanism to Therapeutic Approach" 한국분자세포생물학회 38 (38): 381-389, 2015

62 Huang, J., "Autophagy and human disease" 6 : 1837-1849, 2007

63 Wen, X., "An overview of macroautophagy in yeast" 428 : 1681-1699, 2016

64 Kim, B.W., "ACCORD: an assessment tool to determine the orientation of homodimeric coiledcoils" 7 : 43318-, 2017

65 Liu, X.M., "A selective autophagy pathway takes an unconventional route" 11 : 2381-2382, 2015

66 Mizushima, N., "A protein conjugation system essential for autophagy" 395 : 395-398, 1998

67 Kwon, D.H., "A novel conformation of the LC3-interacting region motif revealed by the structure of a complex between LC3B and RavZ" 490 : 1093-1099, 2017