CD1a, CD1b, and CD1c in immunity against mycobacteria

Advances in Experimental Medicine and Biology

Volumn 783, Issue , 2013, Pages 181-197

  Van Rhijn, Ildiko ^a,b   Ly, Dalam ^a   Moody, D Branch ^a  

^a BRIGHAM AND WOMEN S HOSPITAL   (United States)

^b UTRECHT UNIVERSITY   (Netherlands)

Author keywords

[No Author keywords available]

Indexed keywords

CD1 ANTIGEN; CD1B ANTIGEN; CD1C ANTIGEN; MYCOBACTERIUM ANTIGEN; T6 ANTIGEN; AUTOANTIGEN; BACTERIAL ANTIGEN; CD1C PROTEIN, HUMAN; GLYCOPROTEIN; LIPOPROTEIN; MEMBRANE LIPID;

ANTIGEN BINDING; ANTIGEN EXPRESSION; ANTIGEN FUNCTION; ANTIGEN PRESENTATION; ANTIGEN RECOGNITION; ARTICLE; BACTERIAL IMMUNITY; CELLULAR IMMUNITY; HUMAN; IMMUNOREGULATION; IN VIVO STUDY; INTRACELLULAR TRANSPORT; MOLECULAR MECHANICS; MYCOBACTERIUM; NATURAL KILLER T CELL; NONHUMAN; PRIORITY JOURNAL; T LYMPHOCYTE ACTIVATION; TUBERCULOSIS; ANIMAL; BINDING SITE; CATTLE; CHEMISTRY; ENDOCYTOSIS; ENDOSOME; GUINEA PIG; IMMUNOLOGY; LYSOSOME; METABOLISM; MOUSE; MYCOBACTERIOSIS; PROTEIN TRANSPORT; REVIEW; SPECIES DIFFERENCE; STRUCTURE ACTIVITY RELATION; T LYMPHOCYTE SUBPOPULATION; ULTRASTRUCTURE;

ANIMALS; ANTIGEN PRESENTATION; ANTIGENS, BACTERIAL; ANTIGENS, CD1; AUTOANTIGENS; BINDING SITES; CATTLE; ENDOCYTOSIS; ENDOSOMES; GLYCOPROTEINS; GUINEA PIGS; HUMANS; LIPOPROTEINS; LYSOSOMES; MEMBRANE LIPIDS; MICE; MYCOBACTERIUM; MYCOBACTERIUM INFECTIONS; NATURAL KILLER T-CELLS; PROTEIN TRANSPORT; SPECIES SPECIFICITY; STRUCTURE-ACTIVITY RELATIONSHIP; T-LYMPHOCYTE SUBSETS;

MLCS; MLOWN;

EID: 84879541144     PISSN: 00652598     EISSN: None     Source Type: Book Series    
DOI: 10.1007/978-1-4614-6111-1_10     Document Type: Article

References (79)

1. Calabi F, Jarvis JM, Martin L, Milstein C (1989) Two classes of CD1 genes. Eur J Immunol 19:285-292
2. Roura-Mir C, Wang L, Cheng TY, Matsunaga I, Dascher CC, Peng SL, Fenton MJ, Kirschning C, Moody DB (2005) Mycobacterium Tuberculosis regulates CD1 antigen presentation pathways through TLR-2. J Immunol 175:1758-1766
3. Matsuda JL, Mallevaey T, Scott-Browne J, Gapin L (2008) CD1d-restricted iNKT cells, the 'Swiss-Army knife' of the immune system. Curr Opin Immunol 20:358-368
4. Stronge VS, Salio M, Jones EY, Cerundolo V (2007) A closer look at CD1d molecules: new horizons in studying NKT cells. Trends Immunol 28:455-462
5. Young MH, Gapin L (2011) Group 1 CD1-restricted T cells take center stage. Eur J Immunol 41:592-594
6. Dougan SK, Kaser A, Blumberg RS (2007) CD1 expression on antigen-presenting cells. Curr Top Microbiol Immunol 314:113-141
7. de Jong A, Pena-Cruz V, Cheng TY, Clark RA, van Rhijn I, Moody DB (2010) CD1aautoreactive T cells are a normal component of the human alphabeta T cell repertoire. Nat Immunol 11:1102-1109
8. Dascher CC, Brenner MB (2003) Evolutionary constraints on CD1 structure: insights from comparative genomic analysis. Trends Immunol 24:412-418
9. Jackman RM, Stenger S, Lee A, Moody DB, Rogers RA, Niazi KR, Sugita M, Modlin RL, Peters PJ, Porcelli SA (1998) The tyrosine-containing cytoplasmic tail of CD1b is essential for its efficient presentation of bacterial lipid antigens. Immunity 8:341-351
10. Moody DB, Porcelli SA (2003) Intracellular pathways of CD1 antigen presentation. Nat Rev Immunol 3:11-22
11. Kang SJ, Cresswell P (2002) Regulation of intracellular trafficking of human CD1d by association with MHC class II molecules. EMBO J 21:1650-1660
12. Kaser A, Hava DL, Dougan SK, Chen Z, Zeissig S, Brenner MB, Blumberg RS (2008) Microsomal triglyceride transfer protein regulates endogenous and exogenous antigen presentation by group 1 CD1 molecules. Eur J Immunol 38:2351-2359
13. Moody DB, Briken V, Cheng TY, Roura-Mir C, Guy MR, Geho DH, Tykocinski ML, Besra GS, Porcelli SA (2002) Lipid length controls antigen entry into endosomal and nonendosomal pathways for CD1b presentation. Nat Immunol 3:435-442
14. Sugita M, Grant EP, van Donselaar E, Hsu VW, Rogers RA, Peters PJ, Brenner MB (1999) Separate pathways for antigen presentation by CD1 molecules. Immunity 11:743-752
15. van Rhijn I, Young DC, de Jong A, Vazquez J, Cheng TY, Talekar R, Barral DC, Leon L, Brenner MB, Katz JT, Riese R, Ruprecht RM, O'Connor PB, Costello CE, Porcelli SA, Briken V, Moody DB (2009) CD1c bypasses lysosomes to present a lipopeptide antigen with 12 amino acids. J Exp Med 206:1409-1422
16. Zeissig S, Dougan SK, Barral DC, Junker Y, Chen Z, Kaser A, Ho M, Mandel H, McIntyre A, Kennedy SM, Painter GF, Veerapen N, Besra GS, Cerundolo V, Yue S, Beladi S, Behar SM, Chen X, Gumperz JE, Breckpot K, Raper A, Baer A, Exley MA, Hegele RA, Cuchel M, Rader DJ, Davidson NO, Blumberg RS (2010) Primary deficiency of microsomal triglyceride transfer protein in human abetalipoproteinemia is associated with loss of CD1 function. J Clin Invest 120:2889-2899
17. Porcelli S, Morita CT, Brenner MB (1992) CD1b restricts the response of human CD4-8-T lymphocytes to a microbial antigen. Nature 360:593-597
18. Manolova V, Kistowska M, Paoletti S, Baltariu GM, Bausinger H, Hanau D, Mori L, de Libero G (2006) Functional CD1a is stabilized by exogenous lipids. Eur J Immunol 36: 1083-1092
19. Zajonc DM, Elsliger MA, Teyton L, Wilson IA (2003) Crystal structure of CD1a in complex with a sulfatide self antigen at a resolution of 2.15 A. Nat Immunol 4:808-815
20. Barral DC, Cavallari M, McCormick PJ, Garg S, Magee AI, Bonifacino JS, de Libero G, Brenner MB (2008) CD1a and MHC class I follow a similar endocytic recycling pathway. Traffic 9:1446-1457
21. Madigan CA, Cheng TY, Layre E, Young DC, McConnell MJ, Debono CA, Murry JP, Wei JR, Barry CE 3rd, Rodriguez GM, Matsunaga I, Rubin EJ, Moody DB (2012) Lipidomic discovery of deoxysiderophores reveals a revised mycobactin biosynthesis pathway in Mycobacterium Tuberculosis. Proc Natl Acad Sci U S A 109:1257-1262
22. Moody DB, Young DC, Cheng TY, Rosat JP, Roura-Mir C, O'Connor PB, Zajonc DM, Walz A, Miller MJ, Levery SB, Wilson IA, Costello CE, Brenner MB (2004) T cell activation by lipopeptide antigens. Science 303:527-531
23. de Voss JJ, Rutter K, Schroeder BG, Su H, Zhu Y, Barry CE 3rd (2000) The salicylatederived mycobactin siderophores of Mycobacterium Tuberculosis are essential for growth in macrophages. Proc Natl Acad Sci U S A 97:1252-1257
24. Siegrist MS, Unnikrishnan M, McConnell MJ, Borowsky M, Cheng TY, Siddiqi N, Fortune SM, Moody DB, Rubin EJ (2009) Mycobacterial Esx-3 is required for mycobactin-mediated iron acquisition. Proc Natl Acad Sci U S A 106:18792-18797
25. Zajonc DM, Crispin MD, Bowden TA, Young DC, Cheng TY, Hu J, Costello CE, Rudd PM, Dwek RA, Miller MJ, Brenner MB, Moody DB, Wilson IA (2005) Molecular mechanism of lipopeptide presentation by CD1a. Immunity 22:209-219
26. de Lalla C, Lepore M, Piccolo FM, Rinaldi A, Scelfo A, Garavaglia C, Mori L, de Libero G, Dellabona P, Casorati G (2010) High-frequency and adaptive-like dynamics of human CD1 self-reactive T cells. Eur J Immunol 41:602-610
27. Porcelli S, Brenner MB, Greenstein JL, Balk SP, Terhorst C, Bleicher PA (1989) Recognition of cluster of differentiation 1 antigens by human CD4-CD8-cytolytic T lymphocytes. Nature 341:447-450
28. Vincent MS, Leslie DS, Gumperz JE, Xiong X, Grant EP, Brenner MB (2002) CD1-dependent dendritic cell instruction. Nat Immunol 3:1163-1168
29. Vincent MS, Xiong X, Grant EP, Peng W, Brenner MB (2005) CD1a-, b-, and c-restricted TCRs recognize both self and foreign antigens. J Immunol 175:6344-6351
30. Shamshiev A, Donda A, Carena I, Mori L, Kappos L, de Libero G (1999) Self glycolipids as T-cell autoantigens. Eur J Immunol 29:1667-1675
31. Brigl M, Tatituri RV, Watts GF, Bhowruth V, Leadbetter EA, Barton N, Cohen NR, Hsu FF, Besra GS, Brenner MB (2011) Innate and cytokine-driven signals, rather than microbial antigens, dominate in natural killer T cell activation during microbial infection. J Exp Med 208:1163-1177
32. Batuwangala T, Shepherd D, Gadola SD, Gibson KJ, Zaccai NR, Fersht AR, Besra GS, Cerundolo V, Jones EY (2004) The crystal structure of human CD1b with a bound bacterial glycolipid. J Immunol 172:2382-2388
33. Beckman EM, Porcelli SA, Morita CT, Behar SM, Furlong ST, Brenner MB (1994) Recognition of a lipid antigen by CD1-restricted alpha beta+ T cells. Nature 372:691-694
34. Garcia-Alles LF, Collmann A, Versluis C, Lindner B, Guiard J, Maveyraud L, Huc E, Im JS, Sansano S, Brando T, Julien S, Prandi J, Gilleron M, Porcelli SA, de la Salle H, Heck AJ, Mori L, Puzo G, Mourey L, de Libero G (2011) Structural reorganization of the antigenbinding groove of human CD1b for presentation of mycobacterial sulfoglycolipids. Proc Natl Acad Sci U S A 108:17755-17760
35. Guiard J, Collmann A, Garcia-Alles LF, Mourey L, Brando T, Mori L, Gilleron M, Prandi J, de Libero G, Puzo G (2009) Fatty acyl structures of mycobacterium Tuberculosis sulfoglycolipid govern T cell response. J Immunol 182:7030-7037
36. Layre E, Collmann A, Bastian M, Mariotti S, Czaplicki J, Prandi J, Mori L, Stenger S, de Libero G, Puzo G, Gilleron M (2009) Mycolic acids constitute a scaffold for mycobacterial lipid antigens stimulating CD1-restricted T cells. Chem Biol 16:82-92
37. Moody DB, Reinhold BB, Guy MR, Beckman EM, Frederique DE, Furlong ST, Ye S, Reinhold VN, Sieling PA, Modlin RL, Besra GS, Porcelli SA (1997) Structural requirements for glycolipid antigen recognition by CD1b-restricted T cells. Science 278:283-286
38. Moody DB, Zajonc DM, Wilson IA (2005) Anatomy of CD1-lipid antigen complexes. Nat Rev Immunol 5:387-399
39. Huang S, Cheng TY, Young DC, Layre E, Madigan CA, Shires J, Cerundolo V, Altman JD, Moody DB (2011) Discovery of deoxyceramides and diacylglycerols as CD1b scaffold lipids among diverse groove-blocking lipids of the human CD1 system. Proc Natl Acad Sci U S A 108:19335-19340
40. Gilleron M, Stenger S, Mazorra Z, Wittke F, Mariotti S, Bohmer G, Prandi J, Mori L, Puzo G, de Libero G (2004) Diacylated Sulfoglycolipids are novel mycobacterial antigens stimulating CD1-restricted T Cells during infection with mycobacterium Tuberculosis. J Exp Med 199:649-659
41. de la Salle H, Mariotti S, Angenieux C, Gilleron M, Garcia-Alles LF, Malm D, Berg T, Paoletti S, Maitre B, Mourey L, Salamero J, Cazenave JP, Hanau D, Mori L, Puzo G, de Libero G (2005) Assistance of microbial glycolipid antigen processing by CD1e. Science 310:1321-1324
42. Ernst WA, Maher J, Cho S, Niazi KR, Chatterjee D, Moody DB, Besra GS, Watanabe Y, Jensen PE, Porcelli SA, Kronenberg M, Modlin RL (1998) Molecular interaction of CD1b with lipoglycan antigens. Immunity 8:331-340
43. Gadola SD, Zaccai NR, Harlos K, Shepherd D, Castro-Palomino JC, Ritter G, Schmidt RR, Jones EY, Cerundolo V (2002) Structure of human CD1b with bound ligands at 2.3 A, a maze for alkyl chains. Nat Immunol 3:721-726
44. Garcia-Alles LF, Versluis K, Maveyraud L, Vallina AT, Sansano S, Bello NF, Gober HJ, Guillet V, de la Salle H, Puzo G, Mori L, Heck AJ, de Libero G, Mourey L (2006) Endogenous phosphatidylcholine and a long spacer ligand stabilize the lipid-binding groove of CD1b. EMBO J 25:3684-3692
45. de Libero G, Mori L (2007) Structure and biology of self lipid antigens. Curr Top Microbiol Immunol 314:51-72
46. Sieling PA, Chatterjee D, Porcelli SA, Prigozy TI, Mazzaccaro RJ, Soriano T, Bloom BR, Brenner MB, Kronenberg M, Brennan PJ (1995) CD1-restricted T cell recognition of microbial lipoglycan antigens. Science 269:227-230
47. Relloso M, Cheng TY, Im JS, Parisini E, Roura-Mir C, Debono C, Zajonc DM, Murga LF, Ondrechen MJ, Wilson IA, Porcelli SA, Moody DB (2008) pH-dependent interdomain tethers of CD1b regulate its antigen capture. Immunity 28:774-786
48. Hunger RE, Sieling PA, Ochoa MT, Sugaya M, Burdick AE, Rea TH, Brennan PJ, Belisle JT, Blauvelt A, Porcelli SA, Modlin RL (2004) Langerhans cells utilize CD1a and langerin to efficiently present nonpeptide antigens to T cells. J Clin Invest 113:701-708
49. Prigozy TI, Sieling PA, Clemens D, Stewart PL, Behar SM, Porcelli SA, Brenner MB, Modlin RL, Kronenberg M (1997) The mannose receptor delivers lipoglycan antigens to endosomes for presentation to T cells by CD1b molecules. Immunity 6:187-197
50. Schaible UE, Winau F, Sieling PA, Fischer K, Collins HL, Hagens K, Modlin RL, Brinkmann V, Kaufmann SH (2003) Apoptosis facilitates antigen presentation to T lymphocytes through MHC-I and CD1 in Tuberculosis. Nat Med 9:1039-1046
51. Tailleux L, Schwartz O, Herrmann JL, Pivert E, Jackson M, Amara A, Legres L, Dreher D, Nicod LP, Gluckman JC, Lagrange PH, Gicquel B, Neyrolles O (2003) DC-SIGN is the major Mycobacterium Tuberculosis receptor on human dendritic cells. J Exp Med 197: 121-127
52. van den Elzen P, Garg S, Leon L, Brigl M, Leadbetter EA, Gumperz JE, Dascher CC, Cheng TY, Sacks FM, Illarionov PA, Besra GS, Kent SC, Moody DB, Brenner MB (2005) Apolipoprotein-mediated pathways of lipid antigen presentation. Nature 437:906-910
53. Kasmar AG, van Rhijn I, Cheng TY, Turner M, Seshadri C, Schiefner A, Kalathur RC, Annand JW, de Jong A, Shires J, Leon L, Brenner M, Wilson IA, Altman JD, Moody DB (2011) CD1b tetramers bind {alpha} {beta} T cell receptors to identify a mycobacterial glycolipid-reactive T cell repertoire in humans. J Exp Med 208:1741-1747
54. Montamat-Sicotte DJ, Millington KA, Willcox CR, Hingley-Wilson S, Hackforth S, Innes J, Kon OM, Lammas DA, Minnikin DE, Besra GS, Willcox BE, Lalvani A (2011) A mycolic acid-specific CD1-restricted T cell population contributes to acute and memory immune responses in human Tuberculosis infection. J Clin Invest 121:2493-2503
55. Ulrichs T, Moody DB, Grant E, Kaufmann SH, Porcelli SA (2003) T-cell responses to CD1-presented lipid antigens in humans with Mycobacterium Tuberculosis infection. Infect Immun 71:3076-3087
56. van Rhijn I, Nguyen TK, Michel A, Cooper D, Govaerts M, Cheng TY, van Eden W, Moody DB, Coetzer JA, Rutten V, Koets AP (2009) Low cross-reactivity of T-cell responses against lipids from Mycobacterium bovis and M. avium paraTuberculosis during natural infection. Eur J Immunol 39:3031-3041
57. Spada FM, Grant EP, Peters PJ, Sugita M, Melian A, Leslie DS, Lee HK, van Donselaar E, Hanson DA, Krensky AM, Majdic O, Porcelli SA, Morita CT, Brenner MB (2000) Selfrecognition of CD1 by gamma/delta T cells: implications for innate immunity. J Exp Med 191:937-948
58. Briken V, Jackman RM, Watts GF, Rogers RA, Porcelli SA (2000) Human CD1b and CD1c isoforms survey different intracellular compartments for the presentation of microbial lipid antigens. J Exp Med 192:281-288
59. Sugita M, van Der WN, Rogers RA, Peters PJ, Brenner MB (2000) CD1c molecules broadly survey the endocytic system. Proc Natl Acad Sci USA 97:8445-8450
60. Beckman EM, Melian A, Behar SM, Sieling PA, Chatterjee D, Furlong ST, Matsumoto R, Rosat JP, Modlin RL, Porcelli SA (1996) CD1c restricts responses of mycobacteria-specific T cells. Evidence for antigen presentation by a second member of the human CD1 family. J Immunol 157:2795-2803
61. Moody DB, Ulrichs T, Muhlecker W, Young DC, Gurcha SS, Grant E, Rosat JP, Brenner MB, Costello CE, Besra GS, Porcelli SA (2000) CD1c-mediated T-cell recognition of isoprenoid glycolipids in Mycobacterium Tuberculosis infection. Nature 20(404):884-888
62. Matsunaga I, Bhatt A, Young DC, Cheng TY, Eyles SJ, Besra GS, Briken V, Porcelli SA, Costello CE, Jacobs WR Jr, Moody DB (2004) Mycobacterium Tuberculosis pks12 produces a novel polyketide presented by CD1c to T cells. J Exp Med 200:1559-1569
63. de Jong A, Arce EC, Cheng TY, van Summeren RP, Feringa BL, Dudkin V, Crich D, Matsunaga I, Minnaard AJ, Moody DB (2007) CD1c presentation of synthetic glycolipid antigens with foreign alkyl branching motifs. Chem Biol 14:1232-1242
64. Scharf L, Li NS, Hawk AJ, Garzon D, Zhang T, Fox LM, Kazen AR, Shah S, Haddadian EJ, Gumperz JE, Saghatelian A, Faraldo-Gomez JD, Meredith SC, Piccirilli JA, Adams EJ (2010) The 2.5 a structure of CD1c in complex with a mycobacterial lipid reveals an open groove ideally suited for diverse antigen presentation. Immunity 33:853-862
65. Hiromatsu K, Dascher CC, Leclair KP, Sugita M, Furlong ST, Brenner MB, Porcelli SA (2002) Induction of CD1-restricted immune responses in guinea pigs by immunization with mycobacterial lipid antigens. J Immunol 169:330-339
66. Watanabe Y, Watari E, Matsunaga I, Hiromatsu K, Dascher CC, Kawashima T, Norose Y, Shimizu K, Takahashi H, Yano I, Sugita M (2006) BCG vaccine elicits both T-cell mediated and humoral immune responses directed against mycobacterial lipid components. Vaccine 24:5700-5707
67. Dascher CC, Hiromatsu K, Xiong X, Morehouse C, Watts G, Liu G, McMurray DN, Leclair KP, Porcelli SA, Brenner MB (2003) Immunization with a mycobacterial lipid vaccine improves pulmonary pathology in the guinea pig model of Tuberculosis. Int Immunol 15: 915-925
68. Nguyen TK, Koets AP, Santema WJ, van Eden W, Rutten VP, van Rhijn I (2009) The mycobacterial glycolipid glucose monomycolate induces a memory T cell response comparable to a model protein antigen and no B cell response upon experimental vaccination of cattle. Vaccine 27:4818-4825
69. Felio K, Nguyen H, Dascher CC, Choi HJ, Li S, Zimmer MI, Colmone A, Moody DB, Brenner MB, Wang CR (2009) CD1-restricted adaptive immune responses to Mycobacteria in human group 1 CD1 transgenic mice. J Exp Med 206:2497-2509
70. Lockridge JL, Chen X, Zhou Y, Rajesh D, Roenneburg DA, Hegde S, Gerdts S, Cheng TY, Anderson RJ, Painter GF, Moody DB, Burlingham WJ, Gumperz JE (2011) Analysis of the CD1 antigen presenting system in humanized SCID mice. PLoS ONE 6:e21701
71. Li D, Hong A, Lu Q, Gao GF, Jin B, Screaton GR, Xu XN (2012) A novel role of CD1c in regulating CD1d-mediated NKT cell recognition by competitive binding to Ig-like transcript 4. Int Immunol 24(11):729-737
72. Li D, Wang L, Yu L, Freundt EC, Jin B, Screaton GR, Xu XN (2009) Ig-like transcript 4 inhibits lipid antigen presentation through direct CD1d interaction. J Immunol 182: 1033-1040
73. Ochoa MT, Stenger S, Sieling PA, Thoma-Uszynski S, Sabet S, Cho S, Krensky AM, Rollinghoff M, Nunes Sarno E, Burdick AE, Rea TH, Modlin RL (2001) T-cell release of granulysin contributes to host defense in leprosy. Nat Med 7:174-179
74. Sieling PA, Ochoa MT, Jullien D, Leslie DS, Sabet S, Rosat JP, Burdick AE, Rea TH, Brenner MB, Porcelli SA, Modlin RL (2000) Evidence for human CD4+ T cells in the CD1-restricted repertoire: derivation of mycobacteria-reactive T cells from leprosy lesions. J Immunol 164:4790-4796
75. Stenger S, Mazzaccaro RJ, Uyemura K, Cho S, Barnes PF, Rosat JP, Sette A, Brenner MB, Porcelli SA, Bloom BR, Modlin RL (1997) Differential effects of cytolytic T cell subsets on intracellular infection. Science 276:1684-1687
76. Stenger S, Hanson DA, Teitelbaum R, Dewan P, Niazi KR, Froelich CJ, Ganz T, Thoma-Uszynski S, Melian A, Bogdan C, Porcelli SA, Bloom BR, Krensky AM, Modlin RL (1998) An antimicrobial activity of cytolytic T cells mediated by granulysin. Science 282:121-125
77. Behar SM, Podrebarac TA, Roy CJ, Wang CR, Brenner MB (1999) Diverse TCRs recognize murine CD1. J Immunol 162:161-167
78. Jahng A, Maricic I, Aguilera C, Cardell S, Halder RC, Kumar V (2004) Prevention of autoimmunity by targeting a distinct, noninvariant CD1d-reactive T cell population reactive to sulfatide. J Exp Med 199:947-957
79. van Rhijn I, Young DC, Im JS, Levery SB, Illarionov PA, Besra GS, Porcelli SA, Gumperz J, Cheng TY, Moody DB (2004) CD1d-restricted T cell activation by nonlipidic small molecules. Proc Natl Acad Sci U S A 101:13578-13583