日本組織適合性学会誌第20巻2号

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1 Major Histocompatibility Complex 2013; 20 (2): 平成 25 年度認定 HLA 検査技術者講習会テキスト HLA 1) 1) 1) HLA phla T T TCR phla HLA cognate phla TCR phla HLA TCR phla MHC TCR cognate phla MHC TCR phla MHC 3 1 phla MHC cognate phla MHC TCR molecular mimicry 2 phla MHC cognate phla MHC TCR HLA MHC induced fit 3 TCR cognate phla MHC phla MHC disparate docking TCR phla MHC T phla MHC HLA-IgG B HLA II HLA CD4 T follicular Th-cell phla MHC キーワード : HLA 1. はじめに T pmhc T TCR 1 T HLA TCR HLA T HLA T T TEL:

2 MHC 2013; 20 (2) HLA 図 1 MHC T pmhc T TCR T MHC TCR TCR GVHD Graftversus-host disease T phla T TCR HLA MHC phla MHC 2.T 細胞の直接アロ phla(mhc) 認識と間接アロ HLA(MHC) 認識の違い T phla MHC 2 HLA MHC TCR HLA MHC phla MHC T 2 T pmhc 3 4 T HLA MHC 図 2 T phla pmhc T phla HLA TCR HLA T phla TCR phla HLA MHC T HLA MHC T HLA MHC T CD4 T Th pmhc MHC- IgG 5 phla MHC Th IgG 3.TCR が phla(mhc) を認識する際の一般的な ルール 1) 一般的な TCR と自己 HLA(MHC)/ 非自己ペプチ ド複合体のドッキング様式 T a b 2 TCR phla MHC phla MHC TCR HLA 2 a 110

3 MHC アロ HLA 抗原に対する拒絶反応の基礎免疫学 図3 2013; 20 (2) TCR の可変領域の相補性決定領域 CDR による phla の識別 TCR の a および b 鎖の可変領域のうち phla と接触する部分は相補性決定領域 CDR と呼ばれ TCRa 鎖と b 鎖にそれぞれ 3 つ ずつ存在する A CDRa1 3 は主に phla ホットドック の右半分を CDRb1 3 は主に phla の左半分を認識する B クス構造とペプチド収容溝に またソーセージがペプチ では HLA MHC の 2 つの a ヘリックス構造を青で ドに相当する これを T 細胞は TCRab 鎖の可変領域を ペプチドを赤で示している TCR の各 CDR と接触する 手にたとえると その 3 本の指を用いて識別する 図 phla MHC を 色 分 け し て 示 す と CDR1a 青 3A これらの指は CDR 相補性決定領域 complemen- CDR2a 紫 CDR3a 緑 は phla MHC の主にク tarity determining region 1 2 および 3 と呼ばれており ラス I 分子の a2 ドメインの a ヘリックス またはクラ TCR の a 鎖と b 鎖の可変領域のそれぞれに存在する ス II 分子の b1 ドメインの a ヘリックスとペプチドの N CDR1a CDR2a CDR3a と CDR1b CDR2b 末 端 側 に 結 合 し CDR1b 黄 CDR2b ピ ン ク CDR3b 図 3B CDR3b 水色 はクラス I およびクラス II 分子ともに これまでに明らかにされた TCR/pHLA MHC 複合 phla MHC の主に a1 ドメインの a ヘリックスとペ 体の立体構造解析結果から この識別法は HLA クラス プチドの C 末端側に結合することがわかる すなわち I とクラス II 分子の両方に当てはまる 次のような原則 phla MHC を紙面に示す向きに配置した場合 上述 があることがわかっている の 原 則 は TCR は phla MHC の 中 心 に 配 位 し 1 TCR の a 鎖は 抗原ペプチドの左 N 末端側の 半 TCRa 鎖は phla MHC の左半分を b 鎖は右半分を 分と HLA MHC の下側を認識する 認識する ということになる 2 TCR の b 鎖は 抗原ペプチドの右 C 末端側の 半 TCR と cognate phla MHC との会合は 熱力学的 分と HLA MHC の上側を認識する に安定な典型的な構造をとると考えられるので 多くの 典型的な phla MHC 複合体を真上から すなわ 場合図 4 に示すような立体配置で TCR と phla が会合 ち T 細胞の TCR 側から見た模式図を図 4 に示す ここ すると想定されている それでは TCR とアロ phla 111

4 MHC 2013; 20 (2) HLA 図 4 HLA TCR CDR1 3 HLA MHC 2 a TCRa CDR1a CDR2a CDR3a phla pmhc I a2 a a2 II b1 a b1 N CDR1b CDR2b CDR3b phla pmhc I II a1 a a1 C MHC 4.TCR とアロ phla(mhc) との会合 1) アロ phla(mhc) の形状が cognate phla(mhc) と酷似する場合 (molecular mimicry, 図 5) TCR phla MHC phla MHC TCR HLA-DR1 HA PKYVKQNTLKLAT HA1.7 TCR HLA-DR4 1) MHC I H2-K b dev8 EQYKFYSV 2C TCR H2-K bm3 2) HLA MHC HLA-DR1 DR4 H2-K b H2-K bm3 phla MHC TCR phla MHC HLA-B8 B*08:01 EB FLR FLRGRAYGR LC13 TCR HLA-B44 B*44:05 Macdonald LC13 TCR HLA-B44 B*44:05 EEY EEYLQAFTY LC13 TCR/HLA-B*4405/EEY 3) HLA-B8 B*08:01 HLA-B44 B*44:05 25 FLR EEY cognate phla MHC phla MHC cognate phla MHC TCR phla MHC molecular mimicry TCR 2 TCR phla 112

5 HLA MHC 2013; 20 (2) 図 5 pmhc HLA cognate pmhc HLA moleucular mimicry HLA HLA HLA cognate phla HLA phla TCR phla 2) アロ phla(mhc) の形状が cognate phla(mhc) の形状と異なっていても,TCR のみならずペプチドと HLA(MHC) のいずれもが, 柔軟に構造を変化させて会合する場合 (induced fit) BM3.3 CBA H-2 k CD8 + T TCR MHC I H2-K b pbm1 INFDFNTI VSV8 RGYVYQGL H2- K bm8 pbm8 SQYYYNSL BM3.3 TCR cognate phla MHC BM3.3 TCR 3 pmhc Reiser 4) Auphan- Anezin 5) BM3.3 TCR a CDR3 phla MHC TCR phla MHC 6 A6 HLA-A2 A*02:01 TAX LLF- GYPVYV TCR HLA-A2 A*02:01 Tel1p MLW- GYLQYV Tel1p TAX 2 phla A6 TCR molecular mimicry A6 TCR Tel1p HLA-A2 A*02:01 a2 a A6 TCR TCRb CDR3 6) TCR phla MHC TCR phla TCR HLA 3 induced fit 7 113

6 MHC 2013; 20 (2) アロ HLA 抗原に対する拒絶反応の基礎免疫学 図6 TCR が形状を柔軟に変化させてアロ phla を認識する場合 induced fit アロ HLA を青で示す TCR の一部 赤色で示す の形状が変化することにより 形状が異なるアロ phla にフィットして これを認 識する例を示す 図7 TCR ペプチドおよび HLA のそれぞれが形状を変化させて TCR が多様な phla を認識する場合 induced fit TCR がアロ phla に交差反応する際に TCR ペプチドおよび HLA のそれぞれが一部分 赤色で示す の形状を変化させて 互いに 結合する例を示す この例では 2 つの phla について HLA が同一であり TCR がアロ反応性を示す例ではないことに注意されたい 114

7 HLA MHC 2013; 20 (2) 3) 同一の TCR が cognate phla(mhc) リガンドとの会合とは全く異なるドッキング様式により, アロ phla(mhc) リガンドと会合する場合 (disparate docking) 2C MHC I H2-K b dev8 EQYKFYSV TCR pmhc H2-L d QL9 QLSPFPFDL H2-K b H2-L d 34 dev8 QL9 2C TCR pmhc QL9/H2-L d cognate pmhc dev8/h2-k b 2C TCR a 30 b 15 pmhc CDR 7) TCR pmhc disparate docking 8 5.TCR がアロ phla(mhc) を直接認識するメカニズムに関するまとめ TCR phla MHC T phla MHC cognate phla MHC molecular mimicry TCR 8) TCR phla MHC polyspecificity phla pmhc T 9) TCR phla MHC TCR TCR HLA phla 2C TCR TCR pmhc T 図 8 Cognate phla phla TCR disparate docking HLA HLA TCR cognate phla phla 115

8 MHC 2013; 20 (2) TCR phla MHC cognate phla MHC 6.Th 細胞の間接アロ pmhc 認識は, 移植後の抗ドナー MHC IgG 抗体の産生に必須である T pmhc 1 HLA IgG HLA CD4 + Th B HLA T HLA 9 Conlon CD4 + T MHC 10) H-2 b H-2 d MHC I H2-K b 10A T T B H2-K b 10B B MHC II MHC-II H-2 b H-2 d 図 9 phla B CD4+Th phla HLA phla B phla Th phla HLA HLA Th TCR phla TCR HLA HLA phla 116

9 HLA MHC 2013; 20 (2) 図 10 MHC B MHC-II T A H-2 b H-2 d H2-K b B T H2-K b C B MHC-II H-2 b H2-K b H2-K b 10C B MHC II Th MHC Th pmhc MHC T II I-A b H2-K d T H-2 b H-2 d H2-K b 11A T II I-A bm12 H-2 b H-2 d/bm12 H2-K b 11B Th pmhc B MHC II MHC Th MHC B MHC IgG pmhc IgG 12 1 MHC B MHC MHC B 2 Th2 3 CD4 + Th2 B MHC II MHC 4 Th2 5 Th2 B Th2 B 6 TH2 CD40 CD40L 117

10 MHC 図 ; 20 (2) アロ HLA 抗原に対する拒絶反応の基礎免疫学 抗ドナー MHC 抗体の産生には 直接アロ pmhc 認識ではなく 間接アロ pmhc 認識する Th 細胞のヘルプが重要である A I-Ab/Kd ペプチド複合体に特異的な間接アロ反応性 T 細胞のみを有するマウス H-2b に アロ心臓 H-2d を移植すると抗 H2-Kb 抗体が産生されるが B T 細胞として I-Abm12 を直接アロ認識する T 細胞のみを有するマウス H-2b では 心臓 H-2d/bm12 を移植 しても抗 H2-Kb 抗体は産生されない 図 12 ヘルパー Th2 細胞による B 細胞上に発現する MHC クラス II 分子とアロ MHC 抗原ペプチドの複合体の間接アロ認識により 抗ドナー MHC-IgG 抗体が産生されるしくみ 詳細については テキスト本文を参照のこと 118

11 HLA MHC 2013; 20 (2) 7 B CD40 8 B 9 10 IgM IgG B Th HLA IgG 参考文献 1) Hennecke J, Wiley DC: Structure of a complex of the human alpha/beta T cell receptor (TCR) HA1.7, influenza hemagglutinin peptide, and major histocompatibility complex class II molecule, HLA-DR4 (DRA*0101 and DRB1*0401): insight into TCR cross-restriction and alloreactivity. J Exp Med (195): , ) Luz JG, Huang M, Garcia KC, et al.: Structural comparison of allogeneic and syngeneic T cell receptor-peptide-major histocompatibility complex complexes: a buried alloreactive mutation subtly alters peptide presentation substantially increasing V(beta) Interactions. J Exp Med (195): , ) Macdonald WA, Chen Z, Gras S, et al.: T cell allorecognition via molecular mimicry. Immunity (31): , ) Reiser JB, Darnault C, Gregoire C, et al.: CDR3 loop flexibility contributes to the degeneracy of TCR recognition. Nat Immunol (4): , ) Auphan-Anezin N, Mazza C, Guimezanes A, et al.: Distinct orientation of the alloreactive monoclonal CD8 T cell activation program by three different peptide/mhc complexes. Eur J Immunol (36): , ) Borbulevych OY, Piepenbrink KH, Gloor BE, et al.: T cell receptor cross-reactivity directed by antigen-dependent tuning of peptide-mhc molecular flexibility. Immunity (31): , ) Colf LA, Bankovich AJ, Hanick NA, et al.: How a single T cell receptor recognizes both self and foreign MHC. Cell (129): , ) Gras S, Kjer-Nielsen L, Chen Z, et al.: The structural bases of direct T-cell allorecognition: implications for T-cell-mediated transplant rejection. Immunol Cell Biol (89): , ) Felix NJ, Donermeyer DL, Horvath S, et al.: Alloreactive T cells respond specifically to multiple distinct peptide-mhc complexes. Nat Immunol (8): , ) Conlon TM, Saeb-Parsy K, Cole JL, et al.: Germinal center alloantibody responses are mediated exclusively by indirectpathway CD4 T follicular helper cells. J Immunol (188): ,

12 MHC 2013; 20 (2) HLA Basic immunology of immune response to allogeneic HLA Atsushi Irie 1) and Yasuharu Nishimura 1) 1) Department of Immunogenetics, Graduate School of Medical Sciences, Kumamoto University Alloreactivity of T cells is a main cause of rejection of allografts, and it depends on direct T-cell recognition of and reaction to the allogenic HLA (allo-hla) molecules. If the mechanisms underlying this great paradox in immunology, in which T cells must stringently discriminate non-self peptide from self peptide in a self-hla-restricted manner, while a large number of T cells show allo-hla reactivity, were solved, it would be tremendously beneficial for patients receiving transplantation medicine. Accumulating knowledge for the crystal structures of TCR/peptide/allo-HLA (MHC) complexes has enabled us to compare them to the structures of TCR/cognate ligand, that is, TCR/non-self peptide/self-hla (MHC) complexes. It has been shown that there are at least three patterns of T-cell allo-recognition; 1) molecular mimicry, 2) induced fit and 3) disparate docking. Since not only TCRs but also peptides and HLAs (MHCs) are so flexible to form the three-member complexes, there is no uniform mechanism of T-cell allo-recognition so far and it is yet difficult to predict the magnitude of T cell reactivity to the certain combination of allo-hlas before transplantation. So, the basic mechanisms underlying the T-cell allo-recognition are awaited to be determined by accumulation of more crystal structures of the same TCRs interacting with their cognate ligands and with allo-ligands (peptide/allo-hlas (MHCs)). Finally, it was recently reported that, in murine experiments, the production of anti-donor-hla-specific IgG antibodies in the recipients after transplantation is exclusively mediated by the recipient helper T (Th) cells indirectly recognized allogenic donor MHC-derived peptides presented by the MHC class II molecules expressed on the surface of recipient B cells. The importance of Th cells activated by indirect allo-recognition for the differentiation of B cells into plasma cells producing long-lasting anti-donor-hla-specific IgG antibody that mediates chronic allograft rejection has been elucidated. Key Words: alloreactivity, direct allorecognition, indirect allorecognition, anti-donor HLA antibody

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