{"id":372,"date":"2018-09-21T00:30:51","date_gmt":"2018-09-20T16:30:51","guid":{"rendered":"http:\/\/www.wuchangsong.com\/?p=372"},"modified":"2018-11-19T10:03:48","modified_gmt":"2018-11-19T02:03:48","slug":"%e7%bb%bc%e8%bf%b0%ef%bc%9a%e5%8f%98%e6%b8%a9%e5%8a%a8%e7%89%a9%e7%9a%84%e9%80%82%e5%ba%94%e6%80%a7%e5%85%8d%e7%96%ab","status":"publish","type":"post","link":"http:\/\/www.wuchangsong.com\/?p=372","title":{"rendered":"\u7efc\u8ff0\uff1a\u53d8\u6e29\u52a8\u7269\u7684\u9002\u5e94\u6027\u514d\u75ab"},"content":{"rendered":"<p><strong>Abstract<\/strong><strong>\uff1a<\/strong>\u9002\u5e94\u6027\u514d\u75ab\u7cfb\u7edf\u8d77\u6e90\u4e8e5\u4ebf\u5e74\u524d\u7684\u53d8\u6e29(\u51b7\u8840)\u810a\u690e\u52a8\u7269\u3002\u4f20\u7edf\u4e0a\uff0c\u9002\u5e94\u6027\u514d\u75ab\u7cfb\u7edf\u662f\u7531\u8868\u8fbe\u91cd\u7ec4\u6fc0\u6d3b\u57fa\u56e0(Rag)\u4f9d\u8d56\u6297\u539f\u53d7\u4f53\u7684\u6dcb\u5df4\u7ec6\u80de\u548cMHC\u7ec4\u6210\u7684\u3002\u8fd9\u4e9b\u7279\u5f81\u5b58\u5728\u4e8e\u6240\u6709\u4e0b\u988c\u9aa8\u7684\u810a\u690e\u52a8\u7269\u4e2d\uff0c\u5305\u62ec\u8f6f\u9aa8\u548c\u786c\u9aa8\u9c7c\u3001\u4e24\u6816\u52a8\u7269\u548c\u722c\u884c\u52a8\u7269\uff0c\u53ef\u80fd\u4e5f\u5b58\u5728\u4e8e\u6700\u53e4\u8001\u7684\u4e0b\u989a\u810a\u690e\u52a8\u7269\uff0c\u5373\u706d\u7edd\u7684\u80ce\u76d8\u76ae\u52a8\u7269\u4e2d\u3002\u7136\u800c\uff0c\u968f\u7740\u65e0\u988c\u9c7c\u7c7b\u9002\u5e94\u6027\u514d\u75ab\u7cfb\u7edf\u7684\u53d1\u73b0\uff0c\u57fa\u4e8e\u4e00\u7ec4\u5b8c\u5168\u4e0d\u540c\u7684\u6297\u539f\u53d7\u4f53-\u53ef\u53d8\u6dcb\u5df4\u7ec6\u80de\u53d7\u4f53-t\u548cb\u7ec6\u80de\u7684\u5206\u53c9\uff0c\u4e5f\u8bb8\u662f\u5148\u5929\u6837\u6dcb\u5df4\u7ec6\u80de\uff0c\u53ef\u4ee5\u8ffd\u6eaf\u5230\u6240\u6709\u810a\u690e\u52a8\u7269\u7684\u8d77\u6e90\u3002\u672c\u7efc\u8ff0\u63a2\u8ba8\u6bd4\u8f83\u514d\u75ab\u5b66\u7684\u6700\u65b0\u8fdb\u5c55\uff0c\u4fc3\u8fdb\u6211\u4eec\u5bf9\u9002\u5e94\u6027\u514d\u75ab\u7cfb\u7edf\u7684\u8d77\u6e90\u548c\u529f\u80fd\u7684\u7406\u89e3\u3002<\/p>\n<p><strong>Introduction<\/strong><strong>\uff1a<\/strong>\u5927\u591a\u6570\u5b9a\u4e49\u4e86\u514d\u75ab\u5b66\u9886\u57df\u7684\u7814\u7a76\u90fd\u662f\u5728\u54fa\u4e73\u52a8\u7269\u8eab\u4e0a\u8fdb\u884c\u7684\uff0c\u7279\u522b\u662f\u5728\u8001\u9f20\u548c\u4eba\u7c7b\u8eab\u4e0a\uff0c\u800c\u6211\u4eec\u7684\u5927\u90e8\u5206\u89c4\u5f8b\u548c\u8303\u5f0f\u90fd\u662f\u4ece\u8fd9\u4e9b\u6a21\u578b\u4e2d\u884d\u751f\u51fa\u6765\u7684\u3002\u7136\u800c\uff0cJaneway and Matzinger\u5236\u5b9a\u4e86\u4e00\u6761\u65b0\u7684\u6cd5\u5219\uff0c\u5b83\u8bf4\u660e\u9002\u5e94\u6027\u514d\u75ab\u7684\u884c\u4e3a\u4e0d\u662f\u7531\u5916\u6765\uff0c\u800c\u662f\u901a\u8fc7\u6a21\u5f0f\u8bc6\u522b\u53d7\u4f53(PRRs)\u88ab\u5916\u90e8\u6216\u5185\u90e8\u5371\u9669\u6240\u6fc0\u53d1\u7684\uff0c\u5b83\u8d77\u6e90\u4e8e\u679c\u8747\u9ed1\u8179\u9f20Toll-like receptors\u7684\u7814\u7a76\u3002\u6b64\u5916\uff0c50\u591a\u5e74\u524d\uff0c\u5728\u9e1f\u7c7b\u6cd5\u6c0f\u56ca\u7684\u7814\u7a76\u4e2d\u53d1\u73b0\u4e86\u4e24\u4e2a\u4e3b\u8981\u6dcb\u5df4\u7ec6\u80de\u4e9a\u7fa4(B\u7ec6\u80de\u548cT\u7ec6\u80de)\u7684\u5dee\u5f02\u3002\u4e3a\u4e86\u4e86\u89e3\u9002\u5e94\u6027\u514d\u75ab\u7684\u8d77\u6e90\uff0c\u6211\u4eec\u5fc5\u987b\u4e86\u89e3\u53d8\u6e29\u810a\u690e\u52a8\u7269\uff0c\u4ee5\u53ca\u810a\u690e\u52a8\u7269\u7684\u76f4\u63a5\u7956\u5148\u2014\u8f83\u4f4e\u7b49\u7684\u540e\u53e3\u52a8\u7269\u3002\u5728\u8fc7\u53bb\u7684\u5341\u5e74\u91cc\uff0c\u7a81\u7834\u6027\u7684\u53d1\u73b0\u5e26\u6765\u4e86\u4eba\u4eec\u5bf9\u9002\u5e94\u6027\u514d\u75ab\u8d77\u6e90\u7684\u8d8a\u6765\u8d8a\u591a\u7684\u8ba4\u8bc6\uff0c\u5e76\u63d0\u9192\u6211\u4eec\u6ce8\u610f\u65b0\u7684\u53ef\u80fd\u6027\u3002<\/p>\n<p>\u4e86\u89e3\u514d\u75ab\u8fdb\u5316\u7684\u4e00\u4e2a\u65b9\u4fbf\u7684\u65b9\u6cd5\u662f\u5c06\u514d\u75ab\u7cfb\u7edf\u5212\u5206\u4e3a\u968f\u7740\u65f6\u95f4\u7684\u63a8\u79fb\u800c\u4fdd\u5b88\u7684\u90e8\u5206\uff0c\u800c\u4e0d\u662f\u90a3\u4e9b\u53d8\u5316\u8fc5\u901f\u7684\u90e8\u5206\uff0c\u8fd9\u662fJan Klein\u6700\u521d\u63d0\u51fa\u7684\u60f3\u6cd5\u3002\u67d0\u4e9b\u514d\u75ab\u7279\u5f81\uff0c\u5982\u514d\u75ab\u7403\u86cb\u767dM(IgM)\u7684\u7ed3\u6784\u548c\u529f\u80fd\uff0c\u4ee5\u53ca\u80f8\u817a\u3001\u813e\u810f\u3001\u5e38\u89c4\u03b1\u03b2T\u7ec6\u80de\u53d7\u4f53(TCRs)\u548cMHC\u2161\u7c7b\u5206\u5b50\u7684\u5b58\u5728\uff0c\u5c06\u5728\u672c\u7efc\u8ff0\u4e2d\u8f83\u4e3a\u8be6\u7ec6\u5730\u63cf\u8ff0\uff0c\u5728\u51e0\u4e4e\u6240\u6709\u7684\u6709\u989a\u7c7b\u810a\u690e\u52a8\u7269\uff08gnathostomes\uff09\u4e2d\u90fd\u662f\u9ad8\u5ea6\u4fdd\u5b88\u7684\u3002\u76f8\u53cd\uff0c\u5176\u4ed6\u514d\u75ab\u6210\u5206\uff0c\u5982IgD\u3001\u03b3\u03b4TCR\u3001\u81ea\u7136\u6740\u4f24\u53d7\u4f53(NKRs)\u548c\u975e\u7ecf\u5178\u7684MHC\u5206\u5b50\u90fd\u662f\u53ef\u5851\u6027\u7684\uff0c\u5728\u57fa\u56e0\u6570\u76ee\uff0c\u7ed3\u6784\u57df\u548c\u529f\u80fd\u4e0a\u603b\u662f\u5b58\u5728\u5dee\u5f02\u3002\u8bb0\u4f4f\u8fd9\u4e00\u6807\u51c6\u4e3a\u7406\u89e3\u514d\u75ab\u7cfb\u7edf\u7684\u57fa\u7840\u63d0\u4f9b\u4e86\u4e00\u4e2a\u6846\u67b6\uff1a\u4fdd\u7559\u7ecf\u8fc7\u9a8c\u8bc1\u7684\u4e8b\u5b9e\uff0c\u4f46\u5141\u8bb8\u8fdb\u5316\u4e0a\u7684\u5feb\u901f\u53d8\u5316\u4e0e\u5176\u4ed6\u4e92\u8865\u7279\u5f81\uff0c\u4ee5\u5bf9\u6297\u4e0d\u65ad\u53d8\u5316\u7684\u75c5\u539f\u4f53\uff08\u73af\u5883\uff09\u3002<\/p>\n<p>\u5728\u7814\u7a76\u4efb\u4f55\u7cfb\u7edf\u7684\u8fdb\u5316\u65f6\uff0c\u4eba\u4eec\u90fd\u4f1a\u5408\u7406\u5730\u5047\u8bbe\uff0c\u4e24\u4e2a\u4e0d\u540c\u7269\u79cd\u4e4b\u95f4\u7684\u5171\u540c\u7279\u5f81\u5f88\u53ef\u80fd\u5b58\u5728\u4e8e\u5b83\u4eec\u7684\u5171\u540c\u7956\u5148\u4e2d\u3002\u7136\u800c\uff0c\u4e00\u4e9b\u7c7b\u4f3c\u7684\u7279\u5f81\u53ef\u80fd\u662f\u901a\u8fc7\u8d8b\u540c\u8fdb\u5316\u5f97\u5230\u7684\uff1b\u4f8b\u5982\uff0c\u5728\u65e0\u989a\u548c\u6709\u989a\u7c7b\u810a\u690e\u52a8\u7269\u4e2d\u51fa\u73b0\u4e86\u4e24\u7ec4\u4e0d\u540c\u7684\u6297\u539f\u53d7\u4f53\uff0c\u4ee5\u53ca\u9ca8\u9c7c\u548c\u9a86\u9a7c\u4e2d\u7684\u5355\u57df\u514d\u75ab\u7403\u86cb\u767d\u53ef\u53d8\u533a(V)\u533a\u57df\u7684\u51fa\u73b0\u3002\u6211\u4eec\u4e5f\u5fc5\u987b\u8ba4\u8bc6\u5230\uff0c\u8fd9\u662f\u4e00\u4e2a\u5e38\u89c1\u7684\u4e25\u91cd\u9519\u8bef\uff0c\u5728\u56fe\u4e2d\u7ed8\u5236\u4ee3\u8868\u6027\u7269\u79cd\u7684\u901f\u8bb0\u65b9\u5f0f\u5e76\u4e0d\u610f\u5473\u7740\u4ece\u8f83\u8001\u7684\u5206\u7c7b\u7fa4\u884d\u751f\u51fa\u6765\u7684\u7269\u79cd\u662f\u6700\u8fd1\u5171\u540c\u7956\u5148\u884d\u751f\u7269\u79cd\u7684\u7956\u5148\uff08Fig 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2\uff09\u3002<\/p>\n<p>\u65e0\u988c\u9c7c\u6709T\u7ec6\u80de(both \u03b1\u03b2 and \u03b3\u03b4 T cells) \u548cB\u7ec6\u80de\uff0c\u4ee5\u53ca\u80f8\u817a\u7c7b\u4f3c\u7269\uff0cMHC\u88ab\u8bc1\u660e\u662f\u96be\u4ee5\u627e\u5230\u7684\uff08Fig 1\u30012\uff09\u3002\u5c31\u50cf\u53ef\u53d8\u6dcb\u5df4\u7ec6\u80de\u53d7\u4f53(VLRs)\uff0cMHC\u53ef\u80fd\u662f\u901a\u8fc7\u6536\u655b\u6f14\u5316\u5728\u8fd9\u4e2a\u7fa4\u4f53\u4e2d\u4ea7\u751f\u7684\u3002\u6bcf\u4e00\u7ec4\u4e2d\u7c98\u819c\u9002\u5e94\u6027\u514d\u75ab\u662f\u5b58\u5728\u7684\u548c\u72ec\u7279\u7684\uff0c\u5982\u4e24\u6816\u52a8\u7269\u548c\u786c\u9aa8\u9c7c\u7684\u7814\u7a76(\u548c\u54fa\u4e73\u52a8\u7269)\u3002\u5728\u4e9a\u5206\u7c7b\u7fa4\u4e2d\u53d1\u73b0\u8f6f\u9aa8\u9c7c\u3001\u786c\u9aa8\u9c7c\u548c\u4e24\u6816\u52a8\u7269\u663e\u793a\u51fa\u533a\u522b\u6574\u4e2a\u810a\u690e\u52a8\u7269\u95e8\u7684\u7279\u5f81\u4f46\u4e5f(\u5728\u67d0\u4e9b\u60c5\u51b5\u4e0b)\u6709\u7740\u72ec\u7279\u7684\u7279\u5f81\uff08Fig 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\u2161\u7c7b\u7cfb\u7edf\u7684\u7f3a\u5931\u548c\u67d0\u4e9b\u514d\u75ab\u7403\u86cb\u767d\u4e9a\u578b\u7684\u7f3a\u5931\u3002\u8f6f\u9aa8\u9c7c\u5177\u6709\u72ec\u7279\u7684\u514d\u75ab\u7403\u86cb\u767d\u7ec4\u7ec7\uff0c\u4f7f\u5f97\u6297\u539f\u53d7\u4f53\u57fa\u56e0\u7684\u51fa\u73b0\u5177\u6709\u65b0\u7684\u529f\u80fd\u3002\u867d\u7136NK\u7ec6\u80de\u660e\u663e\u5b58\u5728\u4e8e\u53d8\u6e29\u52a8\u7269\uff0c\u7531\u4e8e\u514d\u75ab\u7cfb\u7edf\u7684\u5feb\u901f\u53d1\u5c55\uff0c\u8bc6\u522bNKRs\u53d8\u5f97\u975e\u5e38\u56f0\u96be\u3002\u7136\u800c\uff0c\u4e0eMHC\u76f8\u5173\u7684NKR\u57fa\u56e0\u5df2\u5728\u6240\u6709\u810a\u690e\u52a8\u7269\u4e2d\u53d1\u73b0\uff0c\u663e\u793a\u51faMHC \u2160\u548cNKRs\u65e9\u671f\u76f8\u5173\u6027\u3002\u5728\u8fc4\u4eca\u7814\u7a76\u7684\u6240\u6709\u9c7c\u7c7b\u548c\u4e24\u6816\u52a8\u7269\u7684MHC\u4e2d\u90fd\u53d1\u73b0\u4e86\u7f16\u7801MHC\u2160\u7c7b\u5206\u5b50\u3001\u514d\u75ab\u86cb\u767d\u9176\u4f53\u548c\u4e0e\u6297\u539f\u52a0\u5de5\u5206\u5b50\u76f8\u5173\u7684\u8f6c\u8fd0\u4f53(TAP)\u57fa\u56e0\u7684\u53e4\u8001\u8c31\u7cfb\uff0c\u4f46\u662f\u5728\u54fa\u4e73\u52a8\u7269\u4e2d\uff0c\u8fd9\u4e2a\u539f\u578b\u5df2\u7ecf\u88ab\u4e00\u4e2a\u5728MHC\u2160\u7c7b\u80bd\u7279\u5f02\u6027\u4e0d\u90a3\u4e48\u4e25\u683c\u7684\u7cfb\u7edf\u6240\u53d6\u4ee3\u3002\u8fdb\u5316\u4e0a\u7684\u4e00\u5927\u98de\u8dc3\u5728\u4e24\u6816\u52a8\u7269\u4e2d\u662f\u660e\u663e\u7684\uff0c\u5b83\u4eec\u8868\u73b0\u51fa\u5178\u578b\u7684\u6297\u4f53\u7c7b\u8f6c\u6362\uff0c\u5e76\u4e14\u6709\u4e00\u4e2aIgG\u7c7b(\u79f0\u4e3aIgY)\uff0c\u5b83\u53c2\u4e0e\u5178\u578b\u7684\u8bb0\u5fc6\u514d\u75ab\u53cd\u5e94\uff08Fig 2\uff09\u3002\u810a\u690e\u52a8\u7269\u8fdb\u5316\u7684\u6700\u540e\u4e00\u4e2a\u91cd\u5927\u8fdb\u5c55\u662f\u6dcb\u5df4\u7ed3\u7684\u51fa\u73b0\u548c\u54fa\u4e73\u52a8\u7269\u751f\u53d1\u4e2d\u5fc3\u7684\u5f62\u6210\uff1b\u53d8\u6e29\u810a\u690e\u52a8\u7269\u5141\u8bb8\u5728\u6ee4\u6ce1\u6811\u7a81\u72b6\u7ec6\u80de(FDCs)\u51fa\u73b0\u4e4b\u524d\u8fdb\u884c\u514d\u75ab\u7814\u7a76\uff0c\u800c\u6811\u7a81\u72b6\u7ec6\u80de\u662f\u54fa\u4e73\u52a8\u7269affinity\u6210\u719f\u7684\u4e3b\u8981\u7ec6\u80de\u3002\u672c\u7efc\u8ff0\u5c06\u6df1\u5165\u7814\u7a76\u9002\u5e94\u6027\u514d\u75ab\u7684\u6240\u6709\u8fd9\u4e9b\u7279\u6027\uff0c\u91cd\u70b9\u8ba8\u8bba\u5f15\u8d77\u6240\u6709\u514d\u75ab\u5b66\u5bb6\u5174\u8da3\u7684\u57fa\u672c\u95ee\u9898\uff0c\u5e76\u63a2\u8ba8\u6211\u4eec\u5728\u672a\u6765\u5341\u5e74\u4e2d\u80fd\u591f\u548c\u5e94\u8be5\u89e3\u51b3\u7684\u95ee\u9898\u3002\u503c\u5f97\u6ce8\u610f\u7684\u662f\uff0c\u6e29\u8840\u6027\u662f\u5728\u4f4e\u7b49\u810a\u690e\u52a8\u7269\u7684\u51e0\u4e2a\u5206\u7c7b\u7fa4\u4e2d\u53d1\u73b0\u7684\uff0c\u4f46\u603b\u7684\u6765\u8bf4\uff0c\u5b83\u4eec\u7684\u9002\u5e94\u6027\u514d\u75ab\u8fd8\u6ca1\u6709\u88ab\u68c0\u6d4b\u3002\u5728\u8fd9\u91cc\uff0c\u6211\u5c06\u96c6\u4e2d\u4e8e\u9c7c\u7c7b\u3001\u4e24\u6816\u7c7b\u548c\u722c\u884c\u52a8\u7269\u65b9\u9762\u7684\u5927\u90e8\u5206\u7814\u7a76\u3002<\/p>\n<p>Fig 1.<\/p>\n<p><img decoding=\"async\" loading=\"lazy\" class=\"alignnone size-full wp-image-373\" src=\"http:\/\/www.wuchangsong.com\/wp-content\/uploads\/2018\/09\/1.png\" alt=\"\" width=\"523\" height=\"568\" \/><\/p>\n<p>Fig 2.<\/p>\n<p><img decoding=\"async\" loading=\"lazy\" class=\"alignnone size-full wp-image-374\" src=\"http:\/\/www.wuchangsong.com\/wp-content\/uploads\/2018\/09\/2.png\" alt=\"\" width=\"848\" height=\"505\" \/><\/p>\n<p><strong>Evolution of antigen receptors<\/strong><\/p>\n<p>\u989a\u53e3\u7c7b\u7684\u6297\u4f53\u548cTCRs\u662f\u514d\u75ab\u7403\u86cb\u767d\u8d85\u5bb6\u65cf\uff08IgSF\uff09\u7684\u6210\u5458\uff0c\u6765\u6e90\u4e8e\u672a\u77e5\u7684\u524d\u4f53\u3002\u63d0\u51fa\u4e86\u4e00\u4e2a\u901a\u7528\u6a21\u578b\uff0c\u5e76\u5728\u4e0b\u9762\u8ba8\u8bba\u3002\u76f8\u6bd4\u4e4b\u4e0b\uff0c\u65e0\u988c\u9c7cVLRs\u5c5e\u4e8e\u53e4\u8001\u7684\u5bcc\u542b\u4eae\u6c28\u9178\u7684\u91cd\u590d\u5e8f\u5217(LRR)\u53d7\u4f53\u5bb6\u65cf\uff0c\u53ef\u80fd\u8d77\u6e90\u4e8e\uff08\u4e5f\u662f\u6700\u76f8\u5173\u7684\u4e00\u79cd\u53d7\u4f53\uff09\u8840\u5c0f\u677f\u4e0a\u8868\u8fbe\u7684\u4e00\u79cd\u7ec6\u80de\u8868\u9762\u53d7\u4f53\u3002\u5728\u6dcb\u5df4\u7ec6\u80de\u53d1\u80b2\u8fc7\u7a0b\u4e2d\uff0c\u4e24\u79cd\u6297\u539f\u53d7\u4f53\u90fd\u53ef\u4ee5\u4ea7\u751f\u5f88\u9ad8\u6c34\u5e73\u7684\u591a\u6837\u6027\uff0c\u4f46\u7528\u4e8e\u4ea7\u751f\u591a\u6837\u6027\u7684\u57fa\u56e0\u91cd\u6392\u673a\u5236\u5219\u5b8c\u5168\u4e0d\u540c\u3002<\/p>\n<p><strong>Variable lymphocyte receptors<\/strong><\/p>\n<p>\u4e4b\u524d\u5bf9\u4f53\u6db2\u514d\u75ab\u548c\u7ec6\u80de\u514d\u75ab\u8fdb\u884c\u7684\u4e00\u4e9b\u7814\u7a76\u8868\u660e\uff0c\u65e0\u989a\u7c7b\u5177\u6709\u9002\u5e94\u6027\u514d\u75ab\uff0c\u5bf9\u4e0d\u540c\u7684\u5916\u6765\u6297\u539f\u548c\u540c\u79cd\u6297\u539f\u6709\u7279\u5f02\u6027\u7684\u5e94\u7b54\u3002\u7136\u800c\uff0c30\u591a\u5e74\u6765\u5bf9\u6297\u4f53\u3001TCRs\u548cMHC\u5206\u5b50\u7684\u5e7f\u6cdb\u7814\u7a76\u88ab\u8bc1\u660e\u662f\u5f92\u52b3\u7684\uff0c\u5927\u591a\u6570\u6bd4\u8f83\u514d\u75ab\u5b66\u5bb6\u90fd\u8ba4\u4e3a\uff0c\u5c3d\u7ba1\u65e9\u671f\u8fdb\u884c\u4e86\u529f\u80fd\u7814\u7a76\uff0c\u4f46\u65e0\u989a\u7c7b\u6ca1\u6709\u9002\u5e94\u6027\u514d\u75ab\u7cfb\u7edf\u5b58\u5728\u3002\u8fd9\u4e00\u5047\u8bbe\u88abPancer\u548cCooper\u8bc1\u660e\u662f\u4e0d\u6b63\u786e\u7684\uff0c\u4ed6\u4eec\u68c0\u67e5\u4e86\u514d\u75ab\u4e86\u7684\u4e03\u9cc3\u9cd7\u4e2d\u6dcb\u5df4\u7ec6\u80de\u7684\u8f6c\u5f55\u7ec4\uff0c\u5e76\u53d1\u73b0\u4e86\u5927\u91cf\u542b\u6709\u4e0d\u540c\u6570\u91cf\u5185\u90e8\u91cd\u590d\u5e8f\u5217\u7684LRR\u86cb\u767d\uff08Fig 3\uff09\u3002\u8fd9\u4e9b\u542bLRR\u7684\u53d7\u4f53\u5728\u6dcb\u5df4\u7ec6\u80de\u4e2a\u4f53\u53d1\u751f\u8fc7\u7a0b\u4e2d\u88ab\u514b\u9686\u8868\u8fbe\u5e76\u901a\u8fc7\u91cd\u6392\u4ea7\u751f\uff0c\u5e76\u88ab\u547d\u540d\u4e3aVLRs\u3002\u7b2c\u4e00\u4e2a\u8981\u7814\u7a76\u7684\u57fa\u56e0\u662fVLRB\uff0c\u8be5\u57fa\u56e0\u5728\u4e00\u7ec4\u4e03\u9cc3\u9cd7\u6dcb\u5df4\u7ec6\u80de\u4e2d\u8868\u8fbe\u3002VLRB\u86cb\u767d\u88ab\u7cd6\u57fa\u78f7\u8102\u9170\u808c\u9187\u951a\u5b9a\u5728\u4e03\u9cc3\u9cd7\u521a\u5f62\u6210\u7684\u6dcb\u5df4\u7ec6\u80de\u8868\u9762\uff0c\u7136\u540e(\u901a\u8fc7\u672a\u77e5\u7684\u673a\u5236)\u5728\u6297\u539f\u523a\u6fc0\u540e\u4ee5\u4e8c\u805a\u4f53\u7684\u4e94\u805a\u4f53\u5f62\u5f0f\u5206\u6ccc\uff08Fig 3\uff09\u3002\u9996\u5148\uff0c\u4eba\u4eec\u8ba4\u4e3a\u523a\u6fc0\u4e03\u9cc3\u9cd7\u6dcb\u5df4\u7ec6\u80de\u7c7b\u4f3c\u4e8e\u54fa\u4e73\u52a8\u7269\u7684\u975eT\u7ec6\u80de\u4f9d\u8d56\u7684\u514d\u75ab\u53cd\u5e94\uff0c\u5373\u6dcb\u5df4\u7ec6\u80de\u4e0a\u6297\u539f\u53d7\u4f53\u7684\u4ea4\u8054\uff0c\u5f88\u53ef\u80fd\u901a\u8fc7PRR\u4e0e\u53e6\u4e00\u4e2a\u4fe1\u53f7\u7ed3\u5408\u5728\u4e00\u8d77\uff0c\u53ef\u80fd\u6fc0\u6d3bB\u7ec6\u80de\u4ee5\u8bf1\u5bfcVLRB\u5206\u6ccc\u3002\u7136\u800c\uff0c\u4ee4\u4eba\u60ca\u8bb6\u7684\u662f\uff0c\u540e\u6765\u53d1\u73b0VLR\u57fa\u56e0\u4f4d\u70b9\u7b2c\u4e8c\u6b21\u91cd\u6392\uff0cVLRA\u57fa\u56e0\u4f4d\u70b9\u7531\u53e6\u4e00\u90e8\u5206\u6dcb\u5df4\u7ec6\u80de\u8868\u8fbe\uff0c\u5176\u8f6c\u5f55\u7ec4\u4e0e\u6709\u989a\u7c7bT\u7ec6\u80de\u76f8\u4f3c\u3002\u8fdb\u4e00\u6b65\u5206\u6790\u53d1\u73b0\uff0c\u7b2c\u4e09\u79cd\u6297\u539f\u53d7\u4f53VLRC\u4e3b\u8981\u5728\u4e0a\u76ae\u548c\u7c98\u819c\u4e2d\u8868\u8fbe\uff0c\u56e0\u6b64\u65e0\u989a\u7c7bT\u7ec6\u80de\u4f3c\u4e4e\u88ab\u5206\u88c2\u6210\u4e24\u4e2a\u4e9a\u7fa4\uff0c\u53ef\u80fd\u7c7b\u4f3c\u6709\u989a\u7c7b\u03b3\u03b4 \u548c \u03b1\u03b2\u7ec6\u80de\u3002<\/p>\n<p>Fig 3.<\/p>\n<p><img decoding=\"async\" loading=\"lazy\" class=\"alignnone size-full wp-image-375\" src=\"http:\/\/www.wuchangsong.com\/wp-content\/uploads\/2018\/09\/3.png\" alt=\"\" width=\"900\" height=\"559\" \/><\/p>\n<p><strong>IgM antibodies<\/strong><\/p>\n<p>\u5728\u6240\u6709\u7684\u6709\u989a\u7c7b\u4e2d\u53d1\u73b0\u591a\u79cd\u6297\u4f53\u4e9a\u578b\u3002\u591a\u5e74\u6765\uff0c\u5c3d\u7ba1\u5728\u6bcf\u4e2a\u810a\u690e\u52a8\u7269\u7c7b\u4e2dIgM\u6709\u5176\u72ec\u7279\u800c\u8010\u4eba\u5bfb\u5473\u7684\u7279\u5f81\uff0cIgM\u4e00\u76f4\u88ab\u8ba4\u4e3a\u662f\u539f\u59cb\u6297\u4f53\u7c7b\u3002\u5728\u54fa\u4e73\u52a8\u7269\u4e2d\uff0cIgM\u662f\u4e00\u79cd\u4e0e\u8fde\u63a5(J)\u94fe\u76f8\u5173\u7684\u4e94\u805a\u4f53\u3002\u8f6f\u9aa8\u9c7c\u4e2d\u7684IgM\u6709\u4e24\u79cd\u5f62\u5f0f\uff1a\u5c31\u50cf\u6240\u6709\u5176\u4ed6\u810a\u690e\u52a8\u7269\u4e00\u6837\uff0c\u5b58\u5728\u7684\u4e00\u79cd\u591a\u91cd\u5f62\u5f0f\uff0c\u4ee5\u53ca\u4e00\u79cd\u5355\u4f53\u5f62\u5f0f\uff0c\u5728\u9ca8\u9c7c\u6297\u539f\u7279\u5f02\u6027\u53cd\u5e94\u4e2d\u6700\u5e38\u89c1\uff0c\u5f88\u53ef\u80fd\u662f\u4ee5T\u7ec6\u80de\u4f9d\u8d56\u7684\u65b9\u5f0f\u4ea7\u751f\u7684\uff08\u8bf7\u6ce8\u610f\uff0c\u867d\u7136\u5728\u6240\u6709\u5176\u4ed6\u810a\u690e\u52a8\u7269\u4e2d\u4e5f\u53d1\u73b0\u4e86\u5355\u805aIGM\uff0c\u4f46\u53ea\u6709\u5728\u8f6f\u9aa8\u9c7c\u4e2d\u624d\u53d1\u73b0\u5b83\u5728\u514d\u75ab\u5e94\u7b54\u8fc7\u7a0b\u4e2d\u5177\u6709\u91cd\u8981\u7684\u751f\u7406\u76f8\u5173\u6027\uff09\u3002\u786c\u9aa8\u9c7c\u7c7bIgM\u7684\u5206\u6ccc\u5f62\u5f0f\u662f\u4e00\u79cd\u56db\u805a\u4f53\uff0c\u800cj\u94fe\u57fa\u56e0\u5728\u8fd9\u4e00\u7ec4\u4e2d\u5df2\u7ecf\u4e22\u5931\uff0c\u56e0\u4e3a\u5b83\u5b58\u5728\u4e8e\u53e4\u8001\u7684\u8f6f\u9aa8\u9c7c\u4e2d\u3002\u6709\u8da3\u7684\u662f\uff0c\u786c\u9aa8\u9c7c\u5206\u6ccc\u7684IgM\u4e2d\u7684\u4e8c\u786b\u952e\u7684\u8303\u56f4\u662f\u5728\u6297\u539f\u7279\u5f02\u6027\u53cd\u5e94\u7684\u8fc7\u7a0b\u4e2d\u4fee\u6539\u7684\uff0c\u8fd9\u79cd\u4eb2\u548c\u529b\u7684\u6210\u719f\u4f3c\u4e4e\u662f\u8fd9\u7c7b\u6297\u4f53\u6240\u72ec\u6709\u7684\u3002\u6b64\u5916\uff0c\u786c\u9aa8\u9c7cIgM\u7684\u8de8\u819c\u5f62\u5f0f\u4e5f\u662f\u4ea4\u66ff\u62fc\u63a5\u7684\uff0c\u56e0\u6b64\u7531\u4e8e\u672a\u77e5\u7684\u529f\u80fd\u539f\u56e0\uff0c\u5b83\u53ea\u6709\u4e09\u4e2a\u4fdd\u5b88(C)\u7ed3\u6784\u57df(\u800c\u4e0d\u662f\u901a\u5e38\u7684\u56db\u4e2a)\u3002\u603b\u4e4b\uff0cIgM\u662f\u786c\u9aa8\u9c7c\u7c7b\u4e2d\u53d1\u73b0\u7684\u4e3b\u8981\u8840\u6e05\u6297\u4f53\uff0c\u662f\u8fd9\u4e9b\u52a8\u7269\u5bf9\u5e38\u89c4\u6297\u539f\u7684\u53cd\u5e94\uff1b\u8f6f\u9aa8\u9c7c\u9664\u4e86\u5728\u9002\u5e94\u6027\u53cd\u5e94\u4e2d\u4f7f\u7528IgM\u4f5c\u4e3a\u5148\u5929\u6297\u4f53\u7c7b\u5916\uff0c\u8fd8\u5728\u9002\u5e94\u6027\u514d\u75ab\u4e2d\u4f7f\u7528\u5176\u4ed6\u6297\u4f53\u4e9a\u578b\uff1b\u4e24\u6816\u7c7bIgM(\u636e\u6211\u4eec\u6240\u77e5)\u7684\u529f\u80fd\u4e0e\u5176\u54fa\u4e73\u52a8\u7269\u7684\u540c\u6e90\u57fa\u56e0\u975e\u5e38\u76f8\u4f3c\u3002<\/p>\n<p><strong>IgD and IgW antibodies<\/strong><\/p>\n<p>IgD\uff0c\u957f\u671f\u4ee5\u6765\u88ab\u8ba4\u4e3a\u662f\u810a\u690e\u52a8\u7269\u4e2d\u65b0\u53d1\u73b0\u7684\u4e00\u79cd\uff0c\u5b9e\u9645\u4e0a\u975e\u5e38\u53e4\u8001\uff0c\u5b83\u53ef\u4ee5\u8ffd\u6eaf\u5230\u6709\u989a\u7c7b\u7684\u8d77\u6e90\uff0c\u4e00\u79cd\u5728\u8f6f\u9aa8\u9c7c\u4e2d\u53d1\u73b0\u7684\u540c\u79cd\u5f02\u578b\uff0c\u53eb\u505aIgW\uff08Fig 3\uff09\u3002\u4e00\u79cd\u7c7b\u4f3c\u4e8eIgD\u7684\u514d\u75ab\u7403\u86cb\u767d\u5728\u786c\u9aa8\u9c7c\u7c7b\u4e2d\u9996\u5148\u88ab\u53d1\u73b0\uff0c1998\u5e74\u4e4b\u524d\uff0cIgD\u53ea\u5728\u4e00\u4e9b\u54fa\u4e73\u52a8\u7269\u4e2d\u88ab\u53d1\u73b0\uff0c\u8fd9\u662f\u975e\u5e38\u4ee4\u4eba\u60ca\u8bb6\u7684\u3002\u968f\u540e\uff0c\u5728\u4e24\u6816\u52a8\u7269\u5c5e\u975e\u6d32\u722a\u87fe\uff08Xenopus\uff09\u57fa\u56e0\u7ec4\u6570\u636e\u5e93\u4e2d\u53d1\u73b0\u4e86IgD\u57fa\u56e0\uff0c\u5e76\u4e0e\u54fa\u4e73\u52a8\u7269\u4e00\u6837\uff0c\u901a\u8fc7IgM\u548cIgD mRNA\u5728\u65b0\u751fB\u7ec6\u80de\u4e2d\u7684\u9009\u62e9\u6027\u526a\u63a5\u6765\u8868\u8fbe\u3002IgD\u3001IgW\u5728\u8f6f\u9aa8\u9c7c\u7c7b\u7684\u540c\u6e90\u6027\u4e5f\u5728\u80ba\u9c7c\uff08lungfish\uff09\u4e2d\u53d1\u73b0\uff0c\u5e76\u4e14\u5728\u8154\u68d8\u9c7c\uff08coelacanths\uff09\u4e2d\uff0c\u5b83\u4f3c\u4e4e\u662f\u4e3b\u8981\u7684\u540c\u5de5\u578b\uff0c\u8fd9\u79cd\u8457\u540d\u7684\u7269\u79cd\u5df2\u7ecf\u5931\u53bb\u4e86IgM\u3002\u6700\u8fd1\u6709\u5173\u786c\u9aa8\u9c7c\u7c7b\u548c\u4eba\u7c7b\u7684\u6570\u636e\u8868\u660e\uff0cIgD\u53c2\u4e0e\u708e\u75c7\u53cd\u5e94\uff0c\u4e0e\u55dc\u78b1\u6027\u7ec6\u80de\u4e0a\u4e00\u4e2a\u672a\u77e5\u7684Fc\u53d7\u4f53\u7ed3\u5408\u3002\u4e0eIgM\u76f8\u6bd4\uff0cIgD\u5728\u7ed3\u6784\u548c\u529f\u80fd\u4e0a\u90fd\u6709\u5f88\u5927\u7684\u53ef\u5851\u6027\uff1bIgM\u5728\u6574\u4e2a\u8fdb\u5316\u8fc7\u7a0b\u4e2d\u662f\u975e\u5e38\u76f8\u4f3c\u7684\uff0c\u800cIgD\u5728C\u7ed3\u6784\u57df\u7684\u6570\u76ee\u4e0a(\u751a\u81f3\u5728IgD\u672c\u8eab\u7684\u5b58\u5728\u4e0b)\u662f\u9ad8\u5ea6\u53ef\u53d8\u7684\uff0c\u5728\u529f\u80fd\u4e0a\u53ef\u80fd\u4e0e\u8de8\u819c(\u5927\u90e8\u5206\u672a\u77e5)\u548c\u5206\u6ccc(\u56fa\u6709\u7ec6\u80de\u6b66\u88c5)\u7684\u5f62\u5f0f\u6709\u5173\u3002<\/p>\n<p><strong>IgG and IgY antibodies<\/strong><\/p>\n<p>\u54fa\u4e73\u52a8\u7269IgG\u5728\u4e24\u6816\u52a8\u7269\u4e2d\u4ee5\u4e00\u79cd\u53eb\u505aIgY\u7684\u540c\u5de5\u578b\u51fa\u73b0\u3002IgY\u4e0e\u514d\u75ab\u7403\u86cb\u767d\u7c7b\u578b\u8f6c\u6362(CSR)\u540c\u65f6\u51fa\u73b0\u3002\u548cIgM\u4e00\u6837\uff0cIgY\u91cd\u94fe\u6709\u56db\u4e2aC\u57df\u3002\u54fa\u4e73\u52a8\u7269IgG\u548cIgE\u5747\u4e0eIgY\u5171\u540c\u7956\u5148\u6709\u5173\uff0cIgE H\u94fe\u7ef4\u63014\u4e2aC\u7ed3\u6784\u57df\uff0c\u800cIgG H\u94fe\u5931\u53bbCH2\u7ed3\u6784\u57df\u3002\u5728\u975e\u6d32\u722a\u87fe\u4e2d\uff0c\u5411IgY\u7684\u8f6c\u6362\u53ca\u5176\u8868\u8fbe\u5b8c\u5168\u4f9d\u8d56\u4e8eT\u7ec6\u80de\uff0c\u800c\u54fa\u4e73\u52a8\u7269\u6240\u5b9a\u4e49\u7684\u5178\u578bB\u7ec6\u80de\u8bb0\u5fc6\u751f\u6210\u5219\u51fa\u73b0\u5728\u8fd9\u4e00\u7fa4\u4f53\u4e2d\u3002\u8de8\u819cIgY\u548cIgG\u7684\u80de\u8d28\u5c3e\u90fd\u6709\u4e00\u4e2a\u65b0\u7684\u4fe1\u53f7\u57fa\u5e8f\uff0c\u5b83\u6709\u52a9\u4e8eb\u7ec6\u80de\u5728\u8bb0\u5fc6\u53cd\u5e94\u4e2d\u7684\u589e\u6b96\u7206\u53d1\u3002\u9664\u4e86T\u7ec6\u80de\u4f9d\u8d56\u6027\u5916\uff0c\u5bf9\u54fa\u4e73\u52a8\u7269\u5916IgY\u7c7b\u8f6c\u6362\uff08\u89c1\u4e0b\u6587\uff09\u7684\u6761\u4ef6\u77e5\u4e4b\u751a\u5c11\u3002\u5728\u70ed\u5e26\u975e\u6d32\u722a\u87fe\uff08Xenopus tropicalis\uff09\u57fa\u56e0\u7ec4\u4e2d\u53d1\u73b0\u4e86\u4e00\u79cd\u4e0eIgY\u76f8\u5173\u4f46\u7f3a\u4e4f\u4e24\u4e2aC-terminal\u7ed3\u6784\u57df\u7684\u540c\u5de5\u578b\uff0c\u79f0\u4e3aIgF\uff0c\u5176\u529f\u80fd\u5c1a\u4e0d\u6e05\u695a\u3002<\/p>\n<p><strong>Other antibody isotypes and light chains<\/strong><\/p>\n<p>\u5728\u4e0d\u540c\u7684\u810a\u690e\u52a8\u7269\u7c7b\u7fa4\u4e2d\u8fd8\u51fa\u73b0\u4e86\u5176\u4ed6\u7684\u201cdead end\u201dH\u94fe\u514d\u75ab\u7403\u86cb\u767d\u4e9a\u578b\uff0c\u6700\u8fd1\u5bf9\u5b83\u8fdb\u884c\u4e86\u8f83\u8be6\u7ec6\u7684\u7efc\u8ff0\uff0c\u5176\u529f\u80fd\u5c1a\u672a\u5f97\u5230\u63a2\u7d22\u3002\u7c98\u819c\u540c\u5de5\u578b\uff0c\u5982\u54fa\u4e73\u52a8\u7269\u7684\u7ecf\u5178IgA\uff0c\u63cf\u8ff0\u5982\u4e0b\u3002<\/p>\n<p>\u4e0e\u591a\u5e74\u6765\u7684\u60f3\u6cd5\u4e0d\u540c\uff0c\u8f7b\u94fe\uff08L\uff09\u5206\u6b67\u4e3a\u03ba \u548c \u03bb\u53ef\u8ffd\u6eaf\u5230\u6709\u989a\u7c7b\u7684\u8d77\u6e90\u3002\u5728\u5927\u591a\u6570\u7684\u53d8\u6e29\u52a8\u7269\u4e2d\uff0c\u6709\u7b2c\u4e09\u6761\u539f\u59cb\u6027L\u94fe\uff0c\u03c3\uff0c\u5b83\u5728\u722c\u884c\u52a8\u7269\u4e2d\u4e22\u5931\u3002\u5728\u67d0\u4e9b\u5206\u7c7b\u7fa4\u4e2d\u5b58\u5728\u7740L\u94fe\u4e9a\u57fa\uff0c\u5b83\u4eec\u4e5f\u662f\u50cfH\u94fe\u4e00\u6837\u7684dead ends\uff0c\u7279\u522b\u662f\u5728\u9c7c\u7c7b\u4e2d\u3002\u5728\u6240\u6709\u8f6f\u9aa8\u9c7c\u4e2d\uff0c\u03bb\u57fa\u56e0\u662f\u2018\u79cd\u7cfb\u8fde\u63a5\u2019\uff0c\u8fd9\u4f7f\u5f97\u6211\u4eec\u80fd\u591f\u5f88\u597d\u5730\u5206\u6790\u9ca8\u9c7c\u4f53\u5185\u6240\u6709\u4e92\u8865\u51b3\u5b9a\u533a(CDRs)\u7684\u4f53\u7ec6\u80de\u9ad8\u7a81\u53d8(SHM)\u3002\u5728\u6240\u6709\u810a\u690e\u52a8\u7269\u4e2d\uff0c\u5bf9\u67d0\u4e9bH\u94fe\u5f02\u6784\u4f53\u7684L\u94fe\u504f\u597d\u5df2\u88ab\u6ce8\u610f\u5230\uff0c\u8fd9\u662f\u591a\u91cdL\u94fe\u7684\u4e00\u4e2a\u6f5c\u5728\u76ee\u7684\u3002\u53e6\u4e00\u79cd\u65b9\u6cd5\u662f\uff0c\u6709\u4eba\u63d0\u51fa\u5f53\u4e0e\u7c7b\u4f3c\u7684H\u94fe\u914d\u5bf9\u65f6\uff0c\u4e0d\u540c\u7684L\u94fe\u5f02\u6784\u4f53\u53ef\u80fd\u5141\u8bb8\u4e0d\u540c\u7684CDR\u6784\u8c61\u3002\u6700\u540e\uff0c\u5982\u679cL\u94fe\u63d0\u4f9b\u4e86\u81ea\u6fc0\u6d3b\u53d7\u4f53\uff0c\u6216\u8005\u7b2c\u4e00\u6761L\u94fe\u57fa\u56e0\u91cd\u6392\u662f\u975e\u751f\u4ea7\u6027\u7684\uff0c\u591a\u91cdL\u94fe\u5141\u8bb8\u5728H\u94fe\u6b63\u5e38\u5de5\u4f5c\u7684\u7ec6\u80de\u4e2d\u8fdb\u884c\u53d7\u4f53\u7f16\u8f91\u3002<\/p>\n<p><strong>\u03b1\u03b2 T cell receptors<\/strong><\/p>\n<p>\u4e00\u822c\u6765\u8bf4\uff0c\u03b1\u03b2tcrs\u5728\u8fdb\u5316\u8fc7\u7a0b\u4e2d\u662f\u975e\u5e38\u4fdd\u5b88\u7684\uff08Fig 1,2\uff09\u3002\u5728\u6280\u672f\u4e0a\u53ef\u884c\u7684\u60c5\u51b5\u4e0b\uff0c\u80da\u80ce\u6216\u65b0\u751f\u513f\u80f8\u817a\u5207\u9664\u5b9e\u9a8c\u8868\u660e\uff0c\u9002\u5e94\u6027\u514d\u75ab\u4f9d\u8d56\u4e8eT\u7ec6\u80de\uff0c\u5305\u62ec\u9ad8\u5bc6\u5ea6\u6297\u4f53\u3001\u6297\u4f53\u8f6c\u6362\u4ee5\u53ca\u5bf9\u540c\u79cd\u5f02\u4f53\u79fb\u690d\u548c\u75c5\u6bd2\u611f\u67d3\u7684\u7ec6\u80de\u514d\u75ab\u3002\u5728\u5df2\u7814\u7a76\u7684\u6240\u6709\u52a8\u7269\u4e2d\uff0cTCR\u591a\u6837\u6027\u76f8\u5f53\u9ad8\uff0c\u9664\u4e86\u5728\u7c7b\u4f3c\u81ea\u7136\u6740\u4f24\u6027T(NKT)\u7ec6\u80de\u7684\u7814\u7a76\u4e2d(\u89c1\u4e0b\u6587)\u3002\u5728\u67d0\u4e9b\u786c\u9aa8\u9c7c\u7c7b\u7684\u6d4b\u5e8f\u4e2d\uff0c\u53d1\u73b0\u5927\u91cf\u7684\u03b1\u03b2 TCRs\u53c2\u4e0e\u4e86\u5b83\u4eec\u5bf9\u6297\u539f(\u5982\u75c5\u6bd2)\u7684\u53cd\u5e94\u3002<\/p>\n<p><strong>\u03b3\u03b4 T cell receptors<\/strong><\/p>\n<p>\u5728TCR\u53d1\u73b0\u7684\u51e0\u5e74\u4e2d\uff0c\u4eba\u4eec\u53d1\u73b0\u4e86\u4e00\u79cd\u65b0\u7684\u91cd\u7ec4\u6297\u539f\u53d7\u4f53\uff0c\u5b83\u6ca1\u6709\u5df2\u77e5\u7684\u529f\u80fd\uff0c\u540e\u6765\u88ab\u547d\u540d\u4e3aTCR\u03b3\u3002\u51e0\u5e74\u540e\u53d1\u73b0\u4e86\u5b83\u7684\u4f19\u4f34TCR\u03b4\uff0c\u03b3\u03b4 T\u7ec6\u80de\u6210\u4e3a\u9002\u5e94\u6027\u514d\u75ab\u7684\u727a\u7272\u54c1\uff0c\u76f4\u5230\u73b0\u5728\u6211\u4eec\u624d\u5f00\u59cb\u5728\u5206\u5b50\u6c34\u5e73\u4e0a\u638c\u63e1\u5b83\u4eec\u5bf9\u6297\u539f\u7684\u8bc6\u522b\uff0c\u81f3\u5c11\u5bf9\u4e8e\u6d89\u53ca\u5929\u7136\u514d\u75ab\u7684\u03b3\u03b4 T\u7ec6\u80de\u6765\u8bf4\u662f\u5982\u6b64\u3002\u5728\u9664\u786c\u9aa8\u9c7c(\u548c\u80ce\u76d8\u54fa\u4e73\u52a8\u7269)\u4ee5\u5916\u7684\u6240\u6709\u53d8\u6e29\u52a8\u7269\u4e2d\uff0c\u90fd\u6709\u5f88\u5927\u6bd4\u4f8b\u7684TCR\u03b4\u57fa\u56e0\u4ee5\u514d\u75ab\u7403\u86cb\u767dV\u533aH\u94fe(IgVH)\u4f5c\u4e3a\u8bc6\u522b\u5143\u4ef6(BOX 1)\u3002\u514d\u75ab\u7403\u86cb\u767dH\u94fe(IgH)\u548cTCR\u03b4\u4f4d\u70b9\u5728\u51e0\u79cd\u810a\u690e\u52a8\u7269\u4e2d\u7684\u8fde\u9501\u63d0\u793a\u8be5\u57fa\u56e0\u6765\u6e90\u4e8e\u987a\u5f0f\u590d\u5236\uff0c\u957f\u671f\u4ee5\u6765\u4eba\u4eec\u90fd\u8ba4\u8bc6\u5230TCRV\u03b4 \u548c IgVH\u5171\u4eab\u5c5e\u6027\uff0c\u7279\u522b\u662f\u5927\u8303\u56f4\u7684CDR3\u5e8f\u5217\u3002IgH\u4e0eTCR\u03b4\u4e4b\u95f4\u7684\u5173\u7cfb\u662f\u53e4\u8001\u7684\uff0c\u901a\u8fc7\u987a\u5f0f\u590d\u5236\uff0c\u5728\u51e0\u79cd\u810a\u690e\u52a8\u7269\u4e2d\u4e00\u76f4\u5b58\u5728\u7740\u5c06IgVH\u5bfc\u5165TCR\u03b4\u4f4d\u70b9\u3002\u6b64\u5916\uff0c\u4ece\u9644\u8fd1\u7684\u514d\u75ab\u7403\u86cb\u767d\u7c07\u4e2d\u53d1\u73b0\u9ca8\u9c7cIgM\u548cIgW V\u7247\u6bb5\u4e0eTCR\u03b4\u591a\u6837\u6027(D)\u548cJ\u7247\u6bb5\u7684\u8f6c\u6362\u662f\u9ad8\u6c34\u5e73\u7684\u3002\u8fd9\u4e9b\u6570\u636e\u8868\u660e\uff0c\u5f88\u5927\u6bd4\u4f8b\u7684\u03b3\u03b4 T\u7ec6\u80de\u4ee5\u4e00\u79cd\u9002\u5e94\u6027\u7684\u65b9\u5f0f\u53d1\u6325\u4f5c\u7528\uff0c\u8fd9\u5728\u54fa\u4e73\u52a8\u7269\u7684\u51e0\u9879\u7814\u7a76\u4e2d\u5f97\u5230\u4e86\u8bc1\u5b9e\u3002\u76ee\u524d\u5c1a\u9700\u8fdb\u4e00\u6b65\u7814\u7a76\u53d8\u6e29\u52a8\u7269\u5916\u80da\u5c42\u4e2d\u03b3\u03b4 T\u7ec6\u80de\u7684\u9002\u5e94\u6027\u53cd\u5e94\uff0c\u8fd9\u4e9b\u5de5\u4f5c\u624d\u521a\u521a\u5f00\u59cb\u3002\u8fd9\u5728\u8f6f\u9aa8\u9c7c\u7c7b\u4e2d\u5c24\u5176\u4ee4\u4eba\u611f\u5174\u8da3\uff0c\u56e0\u4e3aTCR\u03b3 V\u57fa\u56e0\u5df2\u88ab\u660e\u786e\u5730\u8bc1\u660e\u4e3a\u9ad8\u7a81\u53d8\uff0c\u7c7b\u4f3c\u4e8e\u514d\u75ab\u7403\u86cb\u767d\u57fa\u56e0\u4e2d\u68c0\u6d4b\u5230\u7684\u57fa\u56e0\u3002<\/p>\n<p><strong>Generating diversity: AID versus RAG<\/strong><\/p>\n<p>\u91cd\u7ec4\u6fc0\u6d3b\u57fa\u56e0(RAGs)\u7f16\u7801\u7684\u86cb\u767d\u8d28\u88ab\u7528\u6765\u91cd\u6392\u6709\u989a\u7c7b\uff08gnathostomes\uff09\u7684\u6240\u6709\u6297\u539f\u53d7\u4f53\u57fa\u56e0\u3002RAG 1\u4e3b\u8981\u8d1f\u8d23\u5207\u5272\u91cd\u7ec4\u4fe1\u53f7\u5e8f\u5217(RSSs)\uff0c\u5e76\u53c2\u4e0eRSS\u8bc6\u522b\uff0c\u800cRAG2\u5219\u5728\u57fa\u56e0\u91cd\u6392\u8fc7\u7a0b\u4e2d\u5f15\u5bfc\u548c\u914d\u5408RAG1\u3002\u6700\u8fd1\uff0c\u4e00\u79cd\u5305\u62ecRSSs\uff08\u6216\u81f3\u5c11\u672b\u7aef\u5012\u7f6e\u91cd\u590d\uff09\u3001\u63d2\u5165\u4f4d\u70b9\u548c\u57fa\u56e0\u7684\u8f6c\u5ea7\u5b50\u7f16\u7801RAG\u9176\u5728\u4e00\u79cd\u53eb\u505a\u6587\u660c\u9c7c\uff08amphioxus\uff09\u7684\u8f83\u4f4e\u7684\u5b50\u5bab\u5185\u88ab\u53d1\u73b0(\u5148\u4e8e\u810a\u690e\u52a8\u7269)\u3002\u57fa\u56e0\u7ec4\u4e2d\u6709\u51e0\u4e2a\u8f6c\u5ea7\u5b50\uff0c\u7cfb\u7edf\u53d1\u80b2\u5206\u6790\u8868\u660e\uff0c\u8fd9\u4e9b\u8f6c\u5ea7\u5b50\u5728\u8fd9\u4e2a\u7269\u79cd\u548c\u76f8\u5173\u7269\u79cd\u4e2d\u4ecd\u7136\u5f88\u6d3b\u8dc3\u3002\u6b64\u5916\uff0c\u5728\u51e0\u4e2a\u65e0\u810a\u690e\u52a8\u7269\u7269\u79cd\u4e2d\u8fd8\u53d1\u73b0\u4e86\u4e00\u79cd\u79f0\u4e3aTransib\u7684\u8f6c\u5ea7\u5b50\uff0cRAG1\u7684\u50ac\u5316\u6838\u5fc3\uff0c\u8fd9\u8868\u660e\u8fd9\u79cd\u8f6c\u5ea7\u5b50\u662f\u76f8\u5f53\u53e4\u8001\u7684\u3002\u4ecd\u7136\u5b58\u5728\u7684\u95ee\u9898\u662f\uff0cRAG2\u662f\u539f\u59cb\u8f6c\u5ea7\u5b50\u7684\u4e00\u90e8\u5206\uff0c\u8fd8\u662f\u88ab\u62db\u52df\u6765\u534f\u52a9\u57fa\u56e0\u91cd\u6392\u7684\u57fa\u56e0\u7ec4\u4e2d\u7684\u4e00\u4e2a\u73b0\u6709\u57fa\u56e0\uff1f\u8bf7\u6ce8\u610f\uff0c\u8fd9\u4e9bRAGs\u5e76\u4e0d\u7528\u4e8e\u65e0\u988c\u7c7b\uff08agnathans\uff09\u7684VLR\u57fa\u56e0\u91cd\u6392(\u89c1\u4e0b\u6587)\uff0c\u4f46\u5982\u4e0a\u6587\u6240\u8ff0\uff0c\u5728\u5148\u4e8e\u810a\u690e\u52a8\u7269\u7956\u5148\u7684\u540e\u53e3\u52a8\u7269\uff08deuterostomes\uff09\u57fa\u56e0\u7ec4\u4e2d\uff0c\u5df2\u7ecf\u53d1\u73b0\u4e86\u7c7b\u4f3c\u4e8eRAG 1\u548cRAG 2\u7684\u57fa\u56e0\uff0c\u8fd9\u4e9b\u57fa\u56e0\u7684\u529f\u80fd(\u5982\u679c\u6709\u7684\u8bdd)\u662f\u672a\u77e5\u7684\u3002<\/p>\n<p>2000\u5e74\uff0c\u5728\u5c0f\u9f20\u548c\u4eba\u7c7b\u4e2d\u53d1\u73b0\u6fc0\u6d3b\u8bf1\u5bfc\u7684\u80de\u82f7\u8131\u6c28\u9176(AID)\uff0c\u662f\u54fa\u4e73\u52a8\u7269SHM\u548cCSR\u6240\u9700\u7684\u9176\u3002\u4e0d\u4e45\u4e4b\u540e\uff0c\u5728\u6240\u6709\u7684\u6709\u989a\u7c7b\u4e2d\u90fd\u68c0\u6d4b\u5230\u4e86AID\uff0c\u5e76\u5728\u51e0\u4e2a\u7269\u79cd\u4e2d\u663e\u793a\u51fa\u80fd\u591f\u5728\u6b21\u7ea7\u6dcb\u5df4\u7ec4\u7ec7\u4e2d\u8fdb\u884c\u7a81\u53d8\u548c\u8868\u8fbe\u3002SHM\u548cCSR\u5206\u522b\u9700\u8981AID\u6c28\u57fa\u548c\u7fa7\u57fa\u7aef\uff0c\u56e0\u6b64\u786c\u9aa8\u9c7c\u8fd9\u4e00\u660e\u663e\u7f3a\u4e4fCSR\u7684\u7cfb\u7edf\u53d1\u80b2\u7c7b\u7fa4\u7684AID\u80fd\u591f\u4f53\u5916\u8bf1\u5bfcCSR\u4ee4\u4eba\u60ca\u8bb6\u3002\u4e00\u79cd\u53ef\u80fd\u662f\u786c\u9aa8\u9c7c\u4e27\u5931\u4e86CSR\u7684\u80fd\u529b\uff0c\u56e0\u4e3a\u8f6f\u9aa8\u9c7c\u57fa\u56e0\u7684\u7c07\u578b\u7ec4\u7ec7\uff0c\u4ee5\u524d\u88ab\u8ba4\u4e3a\u4e0d\u80fd\u8fdb\u884cCSR\uff08Fig 1\uff09\u3002\u786e\u5b9e\u53ef\u4ee5\u5728\u4e0d\u540c\u7684IgM\u96c6\u7fa4\u4e4b\u95f4\u4ee5\u53caIgW\u548cIgM\u96c6\u7fa4\u4e4b\u95f4\u8fdb\u884cCSR\u3002\u5728\u9ca8\u9c7c\u975e\u7f16\u7801\u533a\u6ca1\u6709\u53d1\u73b0\u5177\u6709\u91cd\u590d\u5143\u4ef6\u6216\u9776\u533a(r=\u817a\u560c\u5464\u6216\u9e1f\u560c\u5464\uff0cg=\u9e1f\u560c\u5464\uff0cy=\u80de\u5627\u5576\u6216\u80f8\u817a\u5627\u5576)\u57fa\u5e8f\u7684\u5178\u578b\u5f00\u5173\u76d2\uff0c\u56e0\u6b64\uff0c\u5207\u6362\u673a\u5236\u548c\u4e0e\u5178\u578bCSR\u7684\u5173\u7cfb\u90fd\u662f\u672a\u77e5\u7684\u3002\u7136\u800c\uff0c\u5728\u6700\u53e4\u8001\u7684\u810a\u690e\u52a8\u7269\u5177\u6709\u5178\u578b\u7684\u9002\u5e94\u6027\u514d\u75ab\u9884\u793a\u7740\u4e00\u4e2a\u65b0\u7684\u7814\u7a76\u9886\u57df\u5173\u4e8eAID \u548c CSR\uff0c\u8fd9\u4e00\u53d1\u73b0\u4ee4\u4eba\u5174\u594b\u3002<\/p>\n<p>AID\u662f\u8f7d\u8102\u86cb\u767dB mRNA\u7f16\u8f91\u9176\u50ac\u5316\u80bd\u6837(APOBEC)\u5bb6\u65cf\u7684\u6210\u5458\u4e4b\u4e00\uff0c\u7b2c\u4e00\u6b21\u88ab\u53d1\u73b0\u662fmRNA\u526a\u63a5\u7684\u4fee\u9970\u5242\u3002\u540e\u6765\uff0c\u5176\u4ed6APOBEC\u6210\u5458\u88ab\u8bc1\u660e\u53c2\u4e0e\u75c5\u6bd2\u9632\u5fa1\u548c\u4fdd\u62a4\u57fa\u56e0\u7ec4\u514d\u53d7\u9006\u8f6c\u5f55\u75c5\u6bd2\u4fb5\u88ad\u3002APOBEC\u5bb6\u65cf\u7684\u4e24\u4e2a\u6210\u5458\u5728\u4e03\u9cc3\u9cd7\u6dcb\u5df4\u7ec6\u80de\u4e2d\u8868\u8fbe\uff0c\u4e00\u4e2a\u5728\u53d1\u80b2\u4e2d\u7684T\u7ec6\u80de\uff08\u80de\u82f7\u8131\u6c28\u91761\uff08CDA1\uff09\uff09\u4e2d\uff0c\u53e6\u4e00\u4e2a\u5728B\u7ec6\u80de\u4e2d\u8868\u8fbe\uff08CDA2\uff09\uff0c\u8fd9\u4e9b\u9176\u53ef\u80fd\u662f\u4ea7\u751fVLR\u591a\u6837\u6027\u6240\u5fc5\u9700\u7684\u3002VLR\u57fa\u56e0\u901a\u8fc7VLR\u76d2\u7684\u540c\u6e90\u6027\u8fde\u63a5\u5728\u4e00\u4e2a\u79f0\u4e3a\u201ccopy choice\u201d\u7684\u8fc7\u7a0b\u4e2d\u7ec4\u88c5\u3002\u663e\u7136\uff0c\u51fa\u73b0\u4e86\u4e24\u79cd\u9176\uff0c\u4e00\u79cd\u7528\u4e8eT\u7ec6\u80de\u7684\u4f53\u7ec6\u80de\u91cd\u6392\uff0c\u53e6\u4e00\u79cd\u7528\u4e8eB\u7ec6\u80de\u7684\u4f53\u7ec6\u80de\u91cd\u6392\uff0c\u5df2\u7ecf\u4e0d\u9700\u8981\u5bf9\u4e00\u79cd\u9176\u590d\u5408\u7269\u8fdb\u884c\u590d\u6742\u7684\u8c03\u63a7\uff0c\u800c\u8fd9\u79cd\u590d\u6742\u7684\u8c03\u63a7\u662f\u4e0e\u5728\u6709\u989a\u7c7b\uff08gnathostome\uff09 T\u7ec6\u80de\u548cB\u7ec6\u80de\u6297\u539f\u53d7\u4f53\u57fa\u56e0\u4e0a\u884c\u4f7f\u529f\u80fd\u7684RAG\u86cb\u767d\u76f8\u5173\u7684\u3002\u8fd9\u4e9bCDA\u9176\u5728\u4f53\u5916\u5df2\u88ab\u8bc1\u660e\u662f\u7a81\u53d8\u4f53\uff0cCDA 1\u6700\u8fd1\u5df2\u88ab\u878d\u5408\u5230\u4e00\u4e2a\u7528\u4e8e\u4f53\u5185\u8bf1\u53d8\u548c\u57fa\u56e0\u6572\u9664\u7684crispr\u76d2\u4e2d\u3002AID\u548c\u76f8\u5173\u5206\u5b50\u7684\u53d1\u73b0\u9884\u793a\u7740\u9002\u5e94\u6027\u514d\u75ab\u3001\u5929\u7136\u514d\u75ab\u548c\u4e00\u822c\u7ec6\u80de\u5185\u5e73\u8861\u7684\u4e00\u6b21\u4ee4\u4eba\u5174\u594b\u7684\u7a81\u7834\u3002\u4f8b\u5982\uff0cAPOBEC\u5bb6\u65cf\u4e5f\u53c2\u4e0e\u4e86\u53cd\u8f6c\u5f55\u5143\u7d20\u7684\u626b\u63cf\uff0c\u5373\u57fa\u56e0\u7ec4\u7684\u4e00\u822c\u4fdd\u62a4\u3002\u8fd9\u4e00\u9886\u57df\u8fd8\u5904\u4e8e\u8d77\u6b65\u9636\u6bb5\uff0c\u901a\u8fc7\u6bd4\u8f83\u7684\u65b9\u6cd5\u6765\u786e\u5b9aAPOBEC\u5bb6\u65cf\u86cb\u767d\u5728\u975e\u5178\u578b\u514d\u75ab\u8fc7\u7a0b\u4e2d\u7684\u4f5c\u7528\u662f\u5f88\u6709\u5e0c\u671b\u7684\u3002<\/p>\n<p><strong>Evolution of the MHC<\/strong><\/p>\n<p>MHC\u5305\u62ec\u7b2c\u4e00\u7c7b\u3001\u7b2c\u4e8c\u7c7b\u548c\u7b2c\u4e09\u7c7b\u533a\u57df\uff0c\u9996\u6b21\u5728\u8f6f\u9aa8\u9c7c\u4e2d\u53d1\u73b0\u3002\u5728\u5927\u591a\u6570\u7269\u79cd\u4e2d\uff0c\u7ecf\u5178\u7684mhc\u2160\u7c7b\u5206\u5b50\u548cmhc\u2161\u7c7b\u5206\u5b50\u7684\u591a\u6001\u6027\u6c34\u5e73\u5f88\u9ad8\u3002\u6b64\u5916\uff0c\u975e\u7ecf\u5178\u7684MHC\u2160\u7c7b\u57fa\u56e0\u4e5f\u5b58\u5728\u4e8e\u6240\u6709\u68c0\u6d4b\u8fc7\u7684\u6709\u989a\u7c7b\u4e2d\uff0c\u901a\u5e38\u4f4d\u4e8e\u4e0eMHC\u4f4d\u70b9\u672c\u8eab\u4e0d\u540c\u7684\u533a\u57df(\u89c1\u4e0b\u6587)\u3002<\/p>\n<p><strong>MHC class II molecules<\/strong><\/p>\n<p>MHC\u2161\u7c7b\u03b1\u548c\u03b2\u94fe\u57fa\u56e0\u5728\u51e0\u4e4e\u6240\u6709\u7684\u6709\u989a\u7c7b\u5747\u6709\u53d1\u73b0\u3002\u901a\u5e38\uff0c\u6709\u4e24\u4e2a\u6216\u4e09\u4e2a\u540c\u578b\uff0c\u5177\u6709\u9ad8\u5ea6\u7684\u591a\u6001\u6027\u3002\u4e0eMHC\u2160\u7c7b\u4e9a\u578b\u76f8\u6bd4\uff0c\u54fa\u4e73\u52a8\u7269MHC\u2161\u7c7b\u4e9a\u578b\u7684\u8fdb\u5316\u8f83\u6162\uff0c\u5728\u53d8\u6e29\u52a8\u7269\u53ef\u68c0\u6d4b\u5230MHC\u2161\u7c7b\u4e9a\u578b\u3002DO\u5206\u5b50\u5728\u54fa\u4e73\u52a8\u7269\u4e2d\u662fII\u7c7b\u86cb\u767d\u8d28\uff0c\u5176\u901a\u8fc7DM\u5206\u5b50\u8c03\u8282\u80bd\u4e0eMHC II\u7c7b\u5206\u5b50\u7684\u7ed3\u5408\uff0c\u5728\u4efb\u4f55\u53d8\u6e29\u52a8\u7269\u4e2d\u5747\u672a\u53d1\u73b0\u3002\u4e8b\u5b9e\u4e0a\uff0cDM\u57fa\u56e0\u9996\u5148\u51fa\u73b0\u5728\u4e24\u6816\u52a8\u7269\u8eab\u4e0a\uff0c\u5728\u786c\u9aa8\u9c7c\u660e\u663e\u7f3a\u4e4f\uff0c\u800c\u4e14\u5230\u76ee\u524d\u4e3a\u6b62\uff0c\u8fd8\u6ca1\u6709\u5728\u8f6f\u9aa8\u9c7c\u4e2d\u53d1\u73b0\u8fc7DM\u57fa\u56e0\u3002\u7814\u7a76\u8fd9\u4e9b\u50ac\u5316\u5242\u7684\u7f3a\u4e4f\u5982\u4f55\u5f71\u54cd\u591a\u80bd\u4e0eMHC\u2161\u7c7b\u5206\u5b50\u7684\u7ed3\u5408\u662f\u5f88\u6709\u610f\u4e49\u7684\uff1b\u5728\u4e24\u6816\u52a8\u7269\u4e2dMHC\u2161\u7c7b\u5206\u5b50\u7684\u751f\u7269\u5316\u5b66\u65b9\u9762\u5df2\u7ecf\u505a\u4e86\u5927\u91cf\u7684\u5de5\u4f5c\uff0c\u4f46\u5728\u786c\u9aa8\u6216\u8f6f\u9aa8\u9c7c\u7c7b\u4e2d\u6240\u505a\u7684\u5de5\u4f5c\u5374\u5f88\u5c11\u3002\u7136\u800c\uff0c\u5728\u6240\u6709\u53d8\u6e29\u52a8\u7269\u4e2d\u90fd\u53d1\u73b0\u4e86\u4e0d\u53d8\u94fe(\u8fd9\u662fMHC\u2161\u7c7b\u5206\u5b50\u7a33\u5b9a\u7ec4\u88c5\u6240\u5fc5\u9700\u7684)\uff0c\u4e0e\u76f8\u5e94\u7684MHC\u2161\u7c7b\u76f8\u5173\u4e0d\u53d8\u94fe\u80bd(CLIP)\u548c\u9884\u671f\u7684\u7ec4\u7ec7\u5206\u5e03\u3002<\/p>\n<p>\u786c\u9aa8\u9c7c\u5df2\u7ecf\u5931\u53bb\u4e86MHC\u2161\u7c7b\u7cfb\u7edf\uff0c\u8fd9\u9996\u5148\u51fa\u73b0\u5728\u9cd5\u9c7c\uff08cod\uff09\u4e2d\uff0c\u540e\u6765\u53c8\u51fa\u73b0\u5728\u5176\u4ed6\u7269\u79cd\u4e2d\u3002\u4eba\u4eec\u65e9\u5c31\u77e5\u9053\uff0c\u514d\u75ab\u540e\u7684\u9cd5\u9c7c\u65e0\u6cd5\u4ea7\u751f\u7279\u5b9a\u7684\u6297\u4f53\u53cd\u5e94\uff0c\u4ece\u672c\u8d28\u4e0a\u8bf4\uff0c\u5bf9\u6bcf\u79cd\u6297\u539f\u4ea7\u751f\u76f8\u540c\u7684IgM\u6297\u4f53\u3002\u751f\u6d3b\u65b9\u5f0f\u5982\u4f55\u5f71\u54cdMHC II\u7c7b\u5206\u5b50\u7684\u7f3a\u4e4f\u5df2\u88ab\u63a8\u6d4b &#8211; \u8fd9\u4e9b\u52a8\u7269\u751f\u6d3b\u5728\u51b7\u6c34\u73af\u5883\u4e2d\uff0c\u4e5f\u8bb8\u7f3a\u4e4f\u75c5\u539f\u4f53\u538b\u529b\u53ef\u80fd\u5bfc\u81f4\u2018use it or lose it\u2019\u7684\u60c5\u51b5\u3002\u6709\u4eba\u63a8\u6d4b\uff0c\u9cd5\u9c7c\u4e2d\u5927\u91cf\u7684\u975e\u7ecf\u5178MHC\u2160\u7c7b\u5206\u5b50\u53ef\u80fd\u4ee5\u67d0\u79cd\u65b9\u5f0f\u5f25\u8865\u4e86MHC\u2161\u7c7b\u5206\u5b50\u7684\u7f3a\u5931(Box1)\uff0c\u4f46\u5176\u4ed6\u542b\u6709MHC\u2161\u7c7b\u5206\u5b50\u7684\u7269\u79cd\u4e5f\u6269\u589e\u4e86MHC\u2160\u7c7b\u57fa\u56e0\u3002<\/p>\n<p><strong>MHC class I molecules<\/strong><\/p>\n<p>\u7ecf\u5178\u7684\u548c\u975e\u7ecf\u5178\u7684MHC\u2160\u7c7b\u5728\u6240\u6709\u7684\u51b7\u8840\u7684\u6709\u989a\u7c7b\u4e2d\u90fd\u6709\u660e\u663e\u7684\u53d1\u73b0\u3002\u7ecf\u5178\u7684I\u7c7b\u5206\u5b50\u88ab\u5176\u9ad8\u6c34\u5e73\u7684\u591a\u6001\u6027\uff0c\u666e\u904d\u5b58\u5728\u7684\u7ec4\u7ec7\u8868\u8fbe\u548c\u5b9a\u4e49\u7684\u80bd\u7ed3\u5408\u6b8b\u57fa\u6240\u8bc6\u522b\uff0c\u8fd9\u4e9b\u6b8b\u57fa\u9501\u5b9a\u5728\u7ed3\u5408\u80bd\u7684\u6c28\u57fa\u548c\u7fa7\u57fa\u672b\u7aef\u3002\u4e0e\u54fa\u4e73\u52a8\u7269\u4e0d\u540c\u7684\u662f\uff0c\u6297\u539f\u5904\u7406\u7684tap\u57fa\u56e0\u548c\u514d\u75ab\u86cb\u767d\u9176\u4f53(\u7279\u522b\u662f\u86cb\u767d\u9176\u4f53\u4e9a\u57fa-\u03b28(PSM\u03b28))\u57fa\u56e0\u4e0eMHC Ia\u57fa\u56e0\u5bc6\u5207\u76f8\u5173\uff0c\u901a\u5e38\u662f\u5728\u5bb6\u7cfb\u4e2d(Box 1)\u3002<\/p>\n<p>Fig 4.<\/p>\n<p><img decoding=\"async\" loading=\"lazy\" class=\"alignnone size-full wp-image-376\" src=\"http:\/\/www.wuchangsong.com\/wp-content\/uploads\/2018\/09\/4.png\" alt=\"\" width=\"720\" height=\"602\" \/><\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p><strong>Lymphoid tissues: evolutionary insights<\/strong><\/p>\n<p><strong>The thymus<\/strong><\/p>\n<p>\u80f8\u817a\u5b58\u5728\u4e8e\u6240\u6709\u7684\u6709\u989a\u7c7b\uff0c\u901a\u5e38\u6709\u5178\u578b\u7684\u76ae\u8d28\u548c\u9ad3\u8d28\u7ec4\u7ec7\u3002\u5b83\u53ef\u4ee5\u4ece\u4e00\u4e2a\u5c0f\u53f6\u5230\u4e00\u4e2a\u591a\u53f6\u751a\u81f3\u4e0d\u8fde\u7eed\u7684\u7ed3\u6784\uff0c\u53d6\u51b3\u4e8e\u6240\u89c2\u5bdf\u7684\u7269\u79cd\u6216\u53d1\u80b2\u9636\u6bb5\u3002\u6b64\u5916\uff0c\u86cb\u767d\u9176\u4f53\u4e9a\u57fa\u03b211(PSM\u03b211; also known as B5T)\u548c\u81ea\u8eab\u514d\u75ab\u8c03\u8282\u5242(AIRE)\u5728\u6709\u989a\u7c7b\u5bb6\u65cf\u65e9\u671f\u51fa\u73b0\uff0c\u8fd9\u8868\u660e\uff0c\u5728\u54fa\u4e73\u52a8\u7269\u548c\u65e9\u671f\u4e0b\u988c\u810a\u690e\u52a8\u7269\u4e2d\uff0c\u6b63\u5411\u548c\u8d1f\u5411\u9009\u62e9\u662f\u4ee5\u7c7b\u4f3c\u7684\u65b9\u5f0f\u53d1\u751f\u7684\uff08Fig 1,2\uff09\u3002\u76f4\u5230\u6700\u8fd1\uff0c\u4eba\u4eec\u8fd8\u76f8\u4fe1\u80f8\u817a\u5728\u65e0\u989a\u7c7b\u4e2d\u662f\u4e0d\u5b58\u5728\u7684\uff0c\u4f46\u968f\u7740\u643a\u5e26VLRA\u7684T\u7ec6\u80de\u7684\u53d1\u73b0\uff0c\u8fd9\u4e2a\u95ee\u9898\u88ab\u91cd\u65b0\u5ba1\u89c6\u3002\u539f\u4f4d\u63a2\u9488\u03b4\u914d\u4f53(for the Notch receptor)\u548c\u53c9\u5934\u76d2\u86cb\u767dN1(FOXN 1)\uff0c\u8fd9\u662f\u4e00\u79cd\u8f6c\u5f55\u56e0\u5b50\uff0c\u88ab\u8bc1\u660e\u662f\u80f8\u817a\u53d1\u80b2\u6240\u5fc5\u9700\u7684\uff0c\u5b83\u5b9a\u4e49\u4e86\u5e7c\u866b\u54bd\u90e8\u7684\u4e00\u79cd\u7ed3\u6784\u3002\u4e0e\u8fd9\u4e9b\u4e0a\u76ae\u7ec6\u80de\u76f8\u5173\u7684\u662f\u6dcb\u5df4\u7ec6\u80de\u8868\u8fbeVLRA\uff08and in later studies, VLRC\uff09\u548cAPOBEC\u5bb6\u65cf\u9176CDA1\uff0c\u5b83\u4eec\u88ab\u8ba4\u4e3a\u662fVLRA\u548cVLRC\u91cd\u6392\u8fc7\u7a0b\u4e2d\u91cd\u8981\u7684\u5206\u5b50\u3002\u5728\u65e0\u989a\u7c7b\u4e2d\u8fd9\u4e00\u7ed3\u6784\u88ab\u547d\u540d\u4e3a\u80f8\u817a\u6837\u4f53\uff0c\u5e76\u88ab\u8ba4\u4e3a\u662f\u80f8\u817a\u7684\u7b49\u4ef7\u7269\u3002\u5230\u76ee\u524d\u4e3a\u6b62\uff0c\u5728\u65e0\u989a\u7c7b\u4e2d\u8fd8\u6ca1\u6709\u53d1\u73b0PSM\u03b211\uff08\u5b9e\u9645\u4e0a\u6ca1\u6709\u4e13\u95e8\u7684\u86cb\u767d\u9176\u4f53\u6210\u5206\uff09\u6216AIRE\u57fa\u56e0\uff0c\u56e0\u6b64\uff0c\u8fdb\u4e00\u6b65\u7684\u7814\u7a76\u4e3a\u5c06T\u7ec6\u80de\u548cB\u7ec6\u80de\u53d1\u80b2\u4e3a\u4e0d\u540c\u7684\u539f\u4ee3\u6dcb\u5df4\u7ec4\u7ec7\u63d0\u4f9b\u6700\u521d\u7684\u7406\u8bba\u57fa\u7840\u3002\u6587\u660c\u9c7c\u7b49\u540e\u53e3\u52a8\u7269\u9cc3\u533a\u7684\u7ed3\u6784\u4e5f\u8868\u8fbeFOXN 1\u548c\u03b4\u914d\u4f53\uff0c\u56e0\u6b64\uff0c\u7814\u7a76\u5b83\u4eec\u5728\u6dcb\u5df4\u7ec6\u80de\u5206\u5316\u4e2d\u7684\u4f5c\u7528(\u5982\u679c\u6709\u7684\u8bdd)\u662f\u5f88\u6709\u610f\u4e49\u7684\u3002<\/p>\n<p><strong>The spleen as the primordial secondary lymphoid organ<\/strong><\/p>\n<p>\u53ea\u6709\u6e29\u8840\u52a8\u7269\u624d\u6709\u6dcb\u5df4\u7ed3\u3001\u4f69\u8036\u6c0f\u6591\u548c\u751f\u53d1\u4e2d\u5fc3\uff0c\u6240\u6709\u8fd9\u4e9b\u90fd\u4f9d\u8d56\u4e8e\u7ec6\u80de\u56e0\u5b50\u6dcb\u5df4\u6bd2\u7d20\u7684\u5f62\u6210\u3002\u7136\u800c\uff0c\u5927\u591a\u6570\u7684\u6709\u989a\u7c7b\uff0c\u5728\u813e\u810f\u4e2d\u786e\u5b9e\u4f1a\u4ea7\u751f\u9002\u5e94\u6027\u7684\u514d\u75ab\u53cd\u5e94\uff0c\u5c06\u6297\u539f\u96c6\u4e2d\u7528\u4e8e\u6297\u539f\u7279\u5f02\u6027T\u7ec6\u80de\u3001B\u7ec6\u80de\u548c\u6297\u539f\u63d0\u5448\u7ec6\u80de(APCs)\u4e4b\u95f4\u7684\u76f8\u4e92\u4f5c\u7528\u3002\u4ee5\u8f6f\u9aa8\u9c7c\u7c7b\u4e3a\u4ee3\u8868\uff0c\u813e\u810f\u53ef\u5206\u4e3a\u7ea2\u9ad3\u548c\u767d\u9ad3\uff0c\u8bb8\u591a\u810a\u690e\u52a8\u7269\u7684B\u7ec6\u80de\u4e5f\u53ef\u5206\u4e3a\u5206\u9694\u533a\uff08\u8bf7\u6ce8\u610f\uff0c\u8fd9\u79cd\u9694\u79bb\u7ed3\u6784\u5df2\u7ecf\u5728\u51e0\u4e2a\u786c\u9aa8\u9c7c\u548c\u4e24\u6816\u52a8\u7269\u5931\u53bb\u4e86\uff09\u3002\u5728\u53d1\u80b2\u8fc7\u7a0b\u4e2d\uff0cB\u7ec6\u80de\u88ab\u8d8b\u5316\u56e0\u5b50CXC-\u8d8b\u5316\u56e0\u5b50\u914d\u4f5313(CXCL 13)\u6240\u5438\u5f15\uff0c\u8be5\u914d\u4f53\u7531\u813e\u810f\u8840\u7ba1\u8868\u8fbe\uff0c\u5f62\u6210\u65b0\u751f\u7684\u767d\u9ad3\u3002\u5728\u54fa\u4e73\u52a8\u7269\u4e2d(\u53ef\u80fd\u8fd8\u6709\u722c\u884c\u52a8\u7269)\uff0cB\u7ec6\u80de\u79fb\u4f4d\u81f3\u6ee4\u6ce1\uff0cT\u7ec6\u80de\u5305\u56f4\u5c0f\u52a8\u8109\u5468\u56f4\u7684\u6dcb\u5df4\u7ba1\u9798\u3002\u5728\u4e24\u6816\u52a8\u7269\u548c\u9c7c\u7c7b\u4e2d\uff0cB\u7ec6\u80de\u4fdd\u6301\u8fd9\u79cd\u80da\u80ce\u7279\u5f81(BOX 1)\u3002\u6709\u8da3\u7684\u662f\uff0c\u5728\u6f14\u5316\u8fc7\u7a0b\u4e2d\uff0cB\u7ec6\u80de\u5728T\u7ec6\u80de\u533a\u5f62\u6210\u4e4b\u524d\u5f62\u6210\u4e86\u5206\u9694\u7ed3\u6784\uff0c\u8fd9\u5728\u7ecf\u5178\u8bc4\u8bba\u6587\u7ae0\u4e2d\u6709\u5e7f\u6cdb\u7684\u8ba8\u8bba\u3002<\/p>\n<p>Fig 5.<\/p>\n<p><img decoding=\"async\" loading=\"lazy\" class=\"alignnone size-full wp-image-377\" src=\"http:\/\/www.wuchangsong.com\/wp-content\/uploads\/2018\/09\/5.png\" alt=\"\" width=\"445\" height=\"319\" \/><\/p>\n<p><strong>When did conventional and follicular dendritic cells emerge?<\/strong><\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p><strong>Mucosal immunity<\/strong><\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p><strong>T helper cell subsets<\/strong><\/p>\n<p>\u54fa\u4e73\u52a8\u7269T\u8f85\u52a9\u7ec6\u80de\u8868\u578b\u5206\u7c7b\u5728\u8fc7\u53bb10\u5e74\u4e2d\u4ece\u7ecf\u5178\u7684Th1\u548cTh2\u7ec6\u80de\u8303\u5f0f\u6269\u5c55\u5230\u5305\u62ecTh17\u7ec6\u80de\u3001T\u6ee4\u6ce1\u8f85\u52a9\u7ec6\u80de(Tfh)\u3001\u8c03\u8282t\u7ec6\u80de(Treg)\u548c\u5176\u4ed6\u51e0\u4e2at\u7ec6\u80de\u4e9a\u7fa4\u3002\u867d\u7136\u8fd8\u9700\u8981\u505a\u5927\u91cf\u7684\u5de5\u4f5c\u6765\u7814\u7a76T\u7ec6\u80de\u5728\u53d8\u6e29\u52a8\u7269\u4e2d\u7684\u529f\u80fd\uff0c\u4f46\u5f88\u53ef\u80fd\u5728\u6240\u6709\u6709\u989a\u7c7b\u4e2d\u90fd\u4f1a\u53d1\u73b0\u8fd9\u6837\u7684\u8868\u578b\uff08Fig 2\uff09\u3002<\/p>\n<p>&nbsp;<\/p>\n<p><strong>Innate-like lymphocytes<\/strong><\/p>\n<p>\u76ee\u524d\u6700\u4ee4\u4eba\u5174\u594b\u7684\u514d\u75ab\u5b66\u7814\u7a76\u9886\u57df\u4e4b\u4e00\u662f\u5bf9\u5148\u5929\u6837\u6dcb\u5df4\u7ec6\u80de\u7684\u7814\u7a76\uff0c\u5982NKT\u7ec6\u80de\u3001\u9ecf\u819c\u76f8\u5173\u4e0d\u53d8T\u7ec6\u80de(MAIT)\u3001B1\u7ec6\u80de\u3001\u8fb9\u7f18\u533aB\u7ec6\u80de\u548c\u5148\u5929\u6dcb\u5df4\u6837\u7ec6\u80de(ILCs)\uff0c\u5305\u62ecNK\u7ec6\u80de\u3002\u5bf9\u5b83\u4eec\u7684\u8fdb\u5316\u7684\u7814\u7a76\u5e94\u8be5\u662f\u4e00\u4e2a\u65b0\u7684\u4f18\u5148\u4e8b\u9879\uff0c\u56e0\u4e3a\u5b83\u8bf1\u4eba\u5730\u63d0\u51fa\uff0cILCs(\u7279\u522b\u662f)\u53ef\u80fd\u65e9\u4e8e\u643a\u5e26\u6297\u539f\u53d7\u4f53\u7684\u6dcb\u5df4\u7ec6\u80de\u51fa\u73b0\u3002<\/p>\n<p>\u5927\u591a\u6570\u514d\u75ab\u5b66\u5bb6\u8ba4\u4e3a\u5148\u5929T\u7ec6\u80de(NKT\u7ec6\u80de\u548cMAIT\u7ec6\u80de)\u662f\u54fa\u4e73\u52a8\u7269\u8fdb\u5316\u540e\u671f\u7684\u8865\u5145\uff0c\u7c7b\u4f3c\u4e8e\u751f\u53d1\u4e2d\u5fc3\uff08Box 2\uff09\u3002\u5f53Robert\u548c\u4ed6\u7684\u540c\u4e8b\u5728\u4e24\u6816\u52a8\u7269\u5c5e\u975e\u6d32\u722a\u87fe\u4e2d\u68c0\u6d4b\u5230NKT\u6837\u7ec6\u80de\u65f6\uff0c\u8fd9\u79cd\u60f3\u6cd5\u5c31\u88ab\u6401\u7f6e\u4e86\uff0c\u8fd9\u4e9b\u7ec6\u80de\u662fXNC\u4e2d\u7684\u4e00\u4e2a\u975e\u7ecf\u5178MHC\u2160\u7c7b\u5206\u5b50\u6240\u7279\u6709\u7684\u3002\u4e0e\u54fa\u4e73\u52a8\u7269NKT\u7ec6\u80de\u4e00\u6837\uff0c\u86d9NKT\u7ec6\u80de\u5177\u6709\u4e0d\u53d8\u6027\u7684TCR\u03b1\u94fe\uff0c\u5e76\u5177\u6709\u6548\u5e94\u7ec6\u80de\u8868\u578b\u3002\u975e\u6d32\u87fe\u870d\u5c5e\u5e7c\u866b\u8868\u8fbe\u4f4e\u6c34\u5e73\u7684\u7ecf\u5178MHC\u2160\u7c7b\u5206\u5b50\uff0c\u521d\u6b65\u8bc1\u636e\u8868\u660e\uff0c\u5e7c\u866bTCR\u5e93\u4e3b\u8981\u7531\u5e26\u6709\u8fd9\u4e9b\u4e0d\u53d8\u4f53TCR\u03b1\u94fe\u7684\u514b\u9686\u7ec4\u6210\u3002\u8fd9\u9879\u5bf9\u4e24\u6816\u52a8\u7269\u7684\u7814\u7a76\uff0c\u4ee5\u53ca\u722c\u884c\u52a8\u7269\u548c\u9e1f\u7c7bCD1\u5206\u5b50\u7684\u9274\u5b9a\uff0c\u6539\u53d8\u4e86NKT\u7684\u51fa\u73b0\u6a21\u5f0f\u3002\u4e0d\u4ec5NKT\u7ec6\u80de\u5728\u8fdb\u5316\u65e9\u671f\u51fa\u73b0\uff0c\u800c\u4e14\u6709\u4eba\u63d0\u51fa\uff0c\u6dcb\u5df4\u7ec6\u80de\u6570\u91cf\u8f83\u5c11\u7684\u52a8\u7269\u4e3b\u8981\u4f7f\u7528\u80fd\u8fc5\u901f\u6fc0\u53d1\u7684T\u7ec6\u80de\uff0c\u5373\u5728\u6297\u539f\u53d7\u4f53\u523a\u6fc0\u540e\u80fd\u8fc5\u901f\u8fdb\u884c\u5230\u6548\u5e94\u5668\u529f\u80fd\uff0c\u800c\u4e14\uff0c\u5e76\u4e0d\u662f\u6240\u6709\u60c5\u51b5\u4e0b\u90fd\u4f7f\u7528\u6709\u5229\u4e8e\u514b\u9686\u9009\u62e9\u7684\u5927\u578bTCR\u8d44\u6e90\u3002Robert\u7684\u7814\u7a76\u8868\u660e\uff0cNKT\u7ec6\u80de\u5c06\u904d\u5e03\u810a\u690e\u52a8\u7269\u4e9a\u95e8\uff0c\u56e0\u4e3a\u5982\u4e0a\u6240\u8ff0\uff0c\u975e\u7ecf\u5178\u7684MHC\u2160\u7c7b\u57fa\u56e0\u8c31\u7cfb\u5b58\u5728\u4e8e\u6240\u6709\u810a\u690e\u52a8\u7269\u4e2d\u3002<\/p>\n<p>NK\u7ec6\u80de\u529f\u80fd\u5df2\u5728\u6240\u6709\u810a\u690e\u52a8\u7269\u79cd\u7c7b\u4e2d\u68c0\u6d4b\u5230\uff0c\u4f46\u5f88\u96be\u8bc6\u522b\u5176\u53d7\u4f53(\u6ce8\u610f\uff0c\u57281990\u5e74\u4ee3\u5bf9\u54fa\u4e73\u52a8\u7269\u7684\u7814\u7a76\u4e2d\u4e5f\u662f\u5982\u6b64)\u3002\u7531\u4e8e\u7075\u957f\u7c7b\u52a8\u7269\u548c\u556e\u9f7f\u7c7b\u52a8\u7269\u751a\u81f3\u4e0d\u4f7f\u7528\u76f8\u540c\u7684\u57fa\u56e0\u5bb6\u65cf\u6765\u7f16\u7801\u5b83\u4eec\u7684\u4e3b\u8981NKRs\uff0c\u6240\u4ee5NKRs\u7684\u8fdb\u5316\u901f\u5ea6\u975e\u5e38\u5feb\uff0c\u8fd9\u4e00\u70b9\u591a\u5e74\u6765\u4e00\u76f4\u5f88\u6e05\u695a\u3002\u8fd9\u79cd\u5feb\u901f\u7684\u8fdb\u5316\u901f\u7387\u4f7f\u5f97\u5728\u53d8\u6e29\u52a8\u7269\u4e2d\u5f88\u96be\u68c0\u6d4b\u5230NKRs\u3002\u5728\u4e24\u6816\u52a8\u7269\u548c\u9c7c\u7c7b\u4e2d\u5df2\u7ecf\u68c0\u6d4b\u5230NKR\u6837IgSF\u86cb\u767d\u7684\u5927\u57fa\u56e0\u5bb6\u65cf\uff0c\u4f46\u5728\u5927\u591a\u6570\u60c5\u51b5\u4e0b\uff0c\u5b83\u4eec\u5728NK\u7ec6\u80de\u8bc6\u522b\u4e2d\u7684\u4f5c\u7528\u672c\u8eab\u8fd8\u6ca1\u6709\u786e\u5b9a\uff0c\u8bb8\u591a\u65e0\u7591\u8fd8\u6709\u5176\u4ed6\u529f\u80fd\u3002\u76f8\u53cd\uff0c\u5f88\u660e\u663e\uff0c\u9c7c\u7c7b\u548c\u4e24\u6816\u52a8\u7269\u4f53\u5185\u6709\u6dcb\u5df4\u7ec6\u80de\uff0c\u5982\u5207\u9664\u80f8\u817a\u7684\u52a8\u7269\u548c\u6ca1\u6709\u643a\u5e26\u4efb\u4f55\u7c7b\u578b\u7684TCRs\u7684\u52a8\u7269\uff0c\u8fd9\u4e9b\u7ec6\u80de\u4ecd\u7136\u80fd\u591f\u8bf1\u5bfc\u7ec6\u80de\u6eb6\u89e3\u3002\u672a\u6765\u7684\u6311\u6218\u5c06\u662f\u5c06\u8fd9\u4e9b\u53d7\u4f53\u4e0e\u7279\u5b9a\u7684\u7ec6\u80de\u529f\u80fd\u8fde\u63a5\u8d77\u6765\u3002<\/p>\n<p>\u5982\u4e0a\u6587\u6240\u8ff0\uff0cMHC\u53ef\u4ee5\u540c\u65f6\u5bb9\u7eb3IgSF\u548cC\u578b\u51dd\u96c6\u7d20\u5bb6\u65cf\u7684NKRs(Fig 4)\uff0c\u6839\u636e\u7814\u7a76\u7684\u5206\u7c7b\u5355\u5143\u800c\u5b9a\u3002\u9e1f\u7c7b\u6709\u4e24\u79cdC\u578b\u51dd\u96c6\u7d20NKRs\uff0c\u5b83\u4eec\u5728MHC\u4e2d\u4e0e\u54fa\u4e73\u52a8\u7269NKRs\u6709\u4eb2\u7f18\u5173\u7cfb(\u751a\u81f3\u662f\u540c\u6e90)\u3002NK\u7ec6\u80dep30\u76f8\u5173\u86cb\u767d(NKp30; also known as NCR3)\u662f\u4e00\u79cd\u7279\u6b8a\u7684IgSF\u6210\u5458\uff0c\u53ef\u6620\u5c04\u5230\u4eba\u7c7b\u7684MHC\uff0c\u662f\u6709\u989a\u7c7b\u6700\u4fdd\u5b88\u7684NKR\uff0c\u4e5f\u5b58\u5728\u4e8e\u8f6f\u9aa8\u9c7c\u7c7b\u4e2d\u3002\u4e24\u6816\u52a8\u7269\u5728MHC\u7684\u2162\u7c7b\u533a\u57df(\u79f0\u4e3aXMIV)\u4e2d\u5177\u6709NKp30\u7684\u76f4\u63a5\u540c\u6e90\u6027\uff0c\u800cNKp30\u5728MHC\u4e4b\u5916\u6613\u4f4d\uff0c\u5e76\u5728\u53e6\u4e00\u4e2a\u67d3\u8272\u4f53\u7684\u7aef\u7c92\u4e0a\u6269\u5c55\u3002NKp30\uff0cB7\u540c\u7cfb\u72696(B7H6; also known as NCR3LG1)\u7684\u914d\u4f53\u4e5f\u5b58\u5728\u4e8e\u8f6f\u9aa8\u9c7c\u4e2d\uff1b\u6709\u8da3\u7684\u662f\uff0c\u5728NKp30\u5df2\u7ecf\u4e22\u5931\u7684\u7269\u79cd\u4e2d\uff0cB7H6\u4e5f\u4e22\u5931\u4e86\uff1b\u76f8\u53cd\uff0c\u5f53NKp30\u57fa\u56e0\u88ab\u6269\u5c55\u65f6\uff0cB7H6\u57fa\u56e0\u4e5f\u88ab\u6269\u5c55\u3002\u4e00\u822c\u6765\u8bf4\uff0c\u9664\u4e86\u4eba\u7c7b\u548c\u556e\u9f7f\u76ee\u52a8\u7269\u4ee5\u5916\u7684\u7269\u79cd\u8fd8\u6ca1\u6709\u5bf9ILCs\u8fdb\u884c\u4efb\u4f55\u8be6\u7ec6\u7684\u68c0\u67e5\u3002\u5df2\u7ecf\u68c0\u6d4b\u5230\u4e0d\u643a\u5e26\u6297\u539f\u53d7\u4f53\u7684\u4e03\u9cc3\u9cd7\u6dcb\u5df4\u7ec6\u80de\uff0c\u5b83\u4eec\u662fNK\u7ec6\u80de\u548c\u5176\u4ed6ILCs\u7684\u5019\u9009\u7ec6\u80de\u3002<\/p>\n<p>Fig 6.<\/p>\n<p><img decoding=\"async\" loading=\"lazy\" class=\"alignnone size-full wp-image-378\" src=\"http:\/\/www.wuchangsong.com\/wp-content\/uploads\/2018\/09\/6.png\" alt=\"\" width=\"654\" height=\"605\" \/><\/p>\n<p><strong>Quo vadis?<\/strong><\/p>\n<p>\u5728\u6bd4\u8f83\u514d\u75ab\u5b66\u9886\u57df\u5fc5\u987b\u89e3\u51b3\u51e0\u4e2a\u4e3b\u8981\u95ee\u9898\uff08Box 3\uff09\uff0c\u8fd9\u91cc\u5df2\u7ecf\u8c08\u5230\u4e86\u8fd9\u4e9b\u95ee\u9898\u3002\u65e0\u988c\u9c7c\u7c7b\u9002\u5e94\u7cfb\u7edf\u7684\u53d1\u73b0\u63d0\u4f9b\u4e86\u8bb8\u591a\u6709\u8da3\u7684\u95ee\u9898\u5305\u62ec\u80f8\u817a\u7684\u539f\u59cb\u4f5c\u7528\uff08\u88ab\u9009\u4e2d\u6216\u9694\u79bb\uff1f\uff09\uff0c\u6dcb\u5df4\u7ec6\u80de\u8d77\u6e90\uff08ILCs\u6216\u6297\u539f\u53d7\u4f53\u643a\u5e26\u7ec6\u80de\uff1f\uff09\uff0cIgSF\u6297\u539f\u53d7\u4f53\u7684\u51fa\u73b0\uff08\u4ee5\u4ec0\u4e48\u5f62\u5f0f\u51fa\u73b0\uff1f\uff09\uff0c\u4ee5\u53ca\u539fMHC\u5728VLRA(T\u7ec6\u80de\u6837)\u6dcb\u5df4\u7ec6\u80de\u4e2d\u7684\u4f5c\u7528\uff08\u8d8b\u540c\uff1f\uff09\u3002\u7279\u522b\u662f\u5bf9\u4e8e\u540e\u8005\uff0c\u5bf9\u7f3a\u4e4f\u7c98\u819c\u6b21\u7ea7\u6dcb\u5df4\u7ec4\u7ec7\u7684\u52a8\u7269\u8fdb\u884c\u8fd9\u4e00\u7cfb\u7edf\u7684\u7814\u7a76\u53ef\u80fd\u4e3a\u7406\u89e3\u9ad8\u5ea6\u590d\u6742\u7684\u54fa\u4e73\u52a8\u7269\u80a0\u76f8\u5173\u6dcb\u5df4\u7ec4\u7ec7\u63d0\u4f9b\u4e00\u4e2a\u6709\u7528\u7684\u6846\u67b6\u3002APOBEC\u5bb6\u65cf\u7684\u8fdb\u5316\u6211\u4eec\u521a\u521a\u7406\u89e3\u70b9\u76ae\u6bdb\u3002\u6beb\u65e0\u7591\u95ee\uff0c\u672a\u6765\u5bf9\u51b7\u8840\u810a\u690e\u52a8\u7269\u514d\u75ab\u7684\u7814\u7a76\u5c06\u4f7f\u6211\u4eec\u611f\u5230\u60ca\u8bb6\uff0c\u66f4\u91cd\u8981\u7684\u662f\uff0c\u6211\u4eec\u5c06\u7ee7\u7eed\u6539\u53d8\u6211\u4eec\u5bf9\u9002\u5e94\u6027\u514d\u75ab\u7cfb\u7edf\u7684\u770b\u6cd5\u3002<\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p>Box 1<\/p>\n<p>Unique adaptive immune features in ectotherms<\/p>\n<ul>\n<li>\u8f6f\u9aa8\u9c7c\u7684\u514d\u75ab\u7403\u86cb\u767d\u91cd\u94fe(H)\u548c\u8f7b\u94fe(L)\u57fa\u56e0\u5b58\u5728\u4e8e\u201ccluster organization\u201d\u4e2d\uff0c\u53d8\u5f02(V)\u3001\u591a\u6837\u6027(D)\u3001\u8fde\u63a5(J)\u548c\u4fdd\u5b88(C)\u5143\u7d20\u5728\u6bcf\u4e2a\u7c07\u4e2d\u5b58\u5728\uff08Fig 3\uff09\u3002\u5c3d\u7ba1\u5982\u6b64\uff0c\u6bcf\u4e2aB\u7ec6\u80de\u90fd\u8868\u73b0\u51fa\u6297\u539f\u53d7\u4f53\u6392\u65a5\uff0c\u5373\u6bcf\u4e2a\u7ec6\u80de\u4ec5\u8868\u8fbe1\u6761H\u94fe\u3002\u8be5\u7ec4\u7ec7\u5141\u8bb8\u4e0d\u540c\u7c7b\u578b\u7684\u514d\u75ab\u7403\u86cb\u767d\u5feb\u901f\u8fdb\u5316\uff0c\u5305\u62ec\u4e00\u79cd\u79f0\u4e3a\u514d\u75ab\u7403\u86cb\u767d\u65b0\u6297\u539f\u53d7\u4f53(IgNAR)\u7684\u5355\u57dfV\u533a\u6297\u539f\u53d7\u4f53\uff0c\u514d\u75ab\u7403\u86cb\u767d\u57fa\u56e0\u4e0eT\u7ec6\u80de\u53d7\u4f53(TCR)\u57fa\u56e0\u7684\u5bc6\u5207\u5173\u8054\uff0c\u4ee5\u53ca\u5df2\u88ab\u201c\u751f\u6b96\u7cfb\u201d\u8fde\u63a5\u8d77\u6765\u7684\u7c07\uff0c\u5e76\u88ab\u8ba4\u4e3a\u5728\u65e9\u671f\u7684\u4e2a\u4f53\u9057\u4f20\u5b66\u548c\u6210\u5e74\u65f6\u671f\u5177\u6709\u72ec\u7279\u7684\u529f\u80fd\u3002<\/li>\n<li>\u786c\u9aa8\u9c7c\u7684\u4e00\u4e2a\u8c31\u7cfb\u5df2\u7ecf\u5931\u53bb\u4e86MHC\u2161\u7c7b\u57fa\u56e0\u3001\u4e0d\u53d8\u94fe\u548cCD4\u5206\u5b50\uff0c\u4e5f\u5c31\u662f\u8bf4\uff0c\u5f88\u660e\u663e\uff0cT\u8f85\u52a9\u7ec6\u80de\u53d1\u80b2\u6240\u5fc5\u9700\u7684\u6240\u6709\u6210\u5206\u90fd\u4e22\u5931\u4e86\u3002\u9cd5\u9c7c\u53ef\u80fd\u901a\u8fc7\u8868\u8fbe\u8fc7\u591a\u7684\u975e\u7ecf\u5178\u7684MHC\u2160\u7c7b\u57fa\u56e0\u6765\u5f25\u8865\uff0c\u53ef\u80fd\u4f1a\u9009\u62e9\u4e00\u4e2a\u5177\u6709\u591a\u4e2a\u81ea\u7136\u6740\u4f24T(NKT)\u6837\u7ec6\u80de\u7684\u7cfb\u7edf\u3002<\/li>\n<li>\u4e24\u6816\u52a8\u7269\u7ecf\u5386\u4e86\u8715\u53d8\uff0c\u5728\u8fd9\u4e00\u8f6c\u53d8\u8fc7\u7a0b\u4e2d\uff0c\u9002\u5e94\u6027\u514d\u75ab\u53d1\u751f\u4e86\u5e7f\u6cdb\u7684\u53d8\u5316\u3002\u672b\u7aef\u8131\u6c27\u6838\u82f7\u9178\u8f6c\u79fb\u9176(TDT)\u5728\u5e7c\u866b\u4e2d\u4e0d\u8868\u8fbe\uff0c\u56e0\u6b64\u514d\u75ab\u7403\u86cb\u767d\u548cTCRs\u4e2d\u7684\u6297\u539f\u53d7\u4f53\u8fde\u63a5\u7f3a\u4e4fN\u533a\uff0c\u56e0\u6b64\u591a\u6837\u6027\u8f83\u4f4e\u3002\u6b64\u5916\uff0c\u5178\u578b\u7684MHC\u2160\u7c7b\u57fa\u56e0\u8868\u8fbe\u4e5f\u8f83\u4f4e\uff0c\u800cMHC\u2161\u7c7b\u57fa\u56e0\u5728\u874c\u86aa(B\u7ec6\u80de\u548c\u6297\u539f\u63d0\u5448\u7ec6\u80de)\u548c\u6210\u4f53(\u6240\u6709\u6dcb\u5df4\u7ec6\u80de)\u4e2d\u7684\u8868\u8fbe\u4e5f\u4e0d\u540c\u3002\u53d8\u6001\u540e\uff0c\u7b2c\u4e8c\u6ce2\u6dcb\u5df4\u7ec6\u80de\u8fc5\u901f\u53d1\u80b2\uff0c\u73b0\u5728\u6709\u5927\u91cf\u4e92\u8865\u51b3\u5b9a\u533a3\uff08CDR3\uff09\u591a\u6837\u6027\u3002\u4e00\u79cd\u6709\u6548\u7684\u5047\u8bbe\u662f\uff0c\u5e7c\u866b\u7684\u4f53\u6db2\u514d\u75ab\u662f\u7531CD4\u7ec6\u80de\u8c03\u8282\u7684\uff0c\u4f46\u514d\u75ab\u7403\u86cb\u767d\u5e93\u7684\u591a\u6837\u6027\u8f83\u4f4e\uff1b\u7ec6\u80de\u514d\u75ab\u53ef\u80fd\u662fNKT\u6837\u7ec6\u80de\u7684\u7ed3\u6784\u57df\u3002\u6291\u5236\u7ec6\u80de\u53ef\u80fd\u5728\u53d8\u6001\u65f6\u51fa\u73b0\uff0c\u4ee5\u9632\u6b62\u5bf9\u65b0\u51fa\u73b0\u7684\u6210\u4f53\u7279\u5f02\u6027\u81ea\u8eab\u6297\u539f\u7684\u81ea\u8eab\u514d\u75ab\u53cd\u5e94\u3002<\/li>\n<li>\u8457\u540d\u7684\u8154\u68d8\u9c7c\u57fa\u56e0\u7ec4\u5e8f\u5217\u63ed\u793a\u4e86\u514d\u75ab\u7403\u86cb\u767dM\uff08IgM\uff09\u7684\u4e27\u5931\uff0c\u662f\u8fc4\u4eca\u4e3a\u6b62\u552f\u4e00\u5177\u6709\u8fd9\u4e00\u7279\u5f81\u7684\u810a\u690e\u52a8\u7269\u7269\u79cd\u3002IgW\u57fa\u56e0\u5ea7\u53ef\u80fd\u5df2\u7ecf\u53d6\u4ee3\u4e86IgM\u7684\u529f\u80fd\uff0c\u4f46\u662f\u529f\u80fd\u7814\u7a76\u5f88\u96be\u8fdb\u884c\uff0c\u56e0\u4e3a\u8fd9\u4e9b\u9c7c\u662f\u7a00\u6709\u548c\/\u6216\u6fd2\u5371\u7269\u79cd\u3002<\/li>\n<li>\u5357\u6781\u9c7c\u7c7b\u514d\u75ab\u7403\u86cb\u767d\u663e\u793a\u51fa\u660e\u663e\u7684\u9009\u62e9\u6027\uff0c\u7279\u522b\u662f\u5728\u5b83\u4eec\u7684\u94f0\u94fe\u533a\u57df\uff0c\u4ee5\u4fbf\u5728\u6781\u51b7\u7684\u6e29\u5ea6\u4e0b\u4fdd\u6301\u5206\u5b50\u7684\u6d3b\u6027\u3002\u4e0e\u5927\u591a\u6570\u54fa\u4e73\u52a8\u7269\u4e0d\u540c\uff0c\u53d8\u6e29\u52a8\u7269\u5177\u6709\u4e0e\u7ecf\u5178\u7684MHC\u2160\u7c7b\u3001\u4e0e\u6297\u539f\u52a0\u5de5\u76f8\u5173\u8f6c\u8fd0\u5b50(TAP)\u548c\u514d\u75ab\u86cb\u767d\u9176\u4f53\u57fa\u56e0\u76f8\u5173\u7684\u8840\u7edf\u3002\u8fd9\u4e9b\u57fa\u56e0\u53c8\u4e0d\u540c\u4e8e\u54fa\u4e73\u52a8\u7269\uff0c\u4e0eMHC II\u7c7b\u57fa\u56e0\u7d27\u5bc6\u76f8\u8fde\u3002\u86cb\u767d\u9176\u4f53\u4e9a\u57fa-\u03b28(PSM\u03b28)\uff0c\u4f46\u4e0d\u5305\u62ecPSM\u03b29\u6216PSM\u03b210\uff0c\u4e0e\u5176\u5728\u514d\u75ab\u86cb\u767d\u9176\u4f53\u7ec4\u7ec7\u4e2d\u7684\u5173\u952e\u529f\u80fd\u76f8\u4e00\u81f4\u3002\u8bf7\u6ce8\u610f\uff0cPSM\u03b28 and PSM\u03b211 (\u80f8\u817a\u86cb\u767d\u9176\u4f53\u7684\u4e00\u90e8\u5206)\u90fd\u662f\u6784\u6210PSM\u03b25\u7684\u540c\u6784\u4f53\uff0c\u8fd9\u518d\u6b21\u8868\u660e\u03b2-\u86cb\u767d\u9176\u4f53\u73af\u7684\u8fd9\u4e00\u6210\u5458\u626e\u6f14\u4e86\u5173\u952e\u89d2\u8272\u3002<\/li>\n<li>\u8f6f\u9aa8\u6027\u9c7c\u7c7b\u3001\u4e24\u6816\u7c7b\u548c\u8154\u68d8\u76ae\u7c7b(\u4ee5\u53ca\u53e4\u8001\u54fa\u4e73\u52a8\u7269\u548c\u9e1f\u7c7b)\u7684TCR\u03b4\u94fe\u5229\u7528\u514d\u75ab\u7403\u86cb\u767dH\u94fe\u7684V\u533a\uff0c\u5176\u57fa\u56e0\u4f4d\u4e8eTCR \u03b1\u03b4\u4f4d\u70b9\u3002\u8fd9\u79cd\u73b0\u8c61\u6700\u65e9\u662f\u5728\u8f6f\u9aa8\u9c7c\u7684IgNAR\u4e2d\u53d1\u73b0\u7684\uff0c\u4e0d\u4e45\u540e\u5728\u888b\u7c7b\u52a8\u7269\u548c\u5176\u4ed6\u7269\u79cd\u4e2d\u88ab\u53d1\u73b0\u4e3a\u4e0e\u03b4\u94fe\u76f8\u5173\u7684\u5355\u4e2aV\u7ed3\u6784\u57df\u6216VH\u7ed3\u6784\u57df\u3002\u4e0e\u03b3\u03b4 TCR\u529f\u80fd\u7684\u7406\u8bba\u4e00\u81f4\uff0c\u8fd9\u4e9b\u53d1\u73b0\u8868\u660e\u8bb8\u591a\u810a\u690e\u52a8\u7269\u7684\u03b3\u03b4 T\u7ec6\u80de\u5177\u6709\u9002\u5e94\u6027\u529f\u80fd\u3002<\/li>\n<li>\u5728\u6240\u6709\u53d7\u68c0\u7684\u53d8\u6e29\u52a8\u7269\u4e2d\uff0cB\u7ec6\u80de\u90fd\u80fd\u541e\u566c\u9897\u7c92\u548c\u5fae\u751f\u7269\u3002\u8fd9\u4e00\u7279\u5f81\u4e5f\u9002\u7528\u4e8e\u54fa\u4e73\u52a8\u7269\u7684B1\u7ec6\u80de\uff0c\u8868\u660e\u53d8\u6e29\u52a8\u7269B\u7ec6\u80de\u548c\u54fa\u4e73\u52a8\u7269B1\u7ec6\u80de\u4e4b\u95f4\u4ee5\u53caB\u7ec6\u80de\u548c\u9ad3\u6837\u7ec6\u80de\u4e4b\u95f4\u5b58\u5728\u7740\u53e4\u8001\u7684\u8054\u7cfb\u3002<\/li>\n<li>\u4e24\u6816\u52a8\u7269\u7684\u6570\u91cf\u6b63\u5728\u51cf\u5c11\uff0c\u6700\u597d\u7684\u7814\u7a76\u53d1\u73b0\u4e86\u5148\u5929\u514d\u75ab\u673a\u5236\u548c\u75c5\u539f\u4f53\u5bf9\u5b83\u4eec\u7684\u6291\u5236\u4f5c\u7528\u3002\u7136\u800c\uff0c\u4e00\u4e9b\u7814\u7a76\u4e5f\u6697\u793a\u4e86\u9002\u5e94\u6027\u514d\u75ab\u673a\u5236\uff0c\u7279\u522b\u662fMHC\u591a\u6001\u6027\uff0c\u8fd9\u4e9b\u673a\u5236\u4e0e\u4e73\u7cdc\u83cc\u7684\u6613\u611f\u6027\u6216\u6297\u836f\u6027\u6709\u5173\u3002<\/li>\n<li>\u8bb8\u591a\u9c7c\u7c7b\u548c\u4e24\u6816\u52a8\u7269\u90fd\u662f\u591a\u500d\u4f53\u7684\uff0c\u9002\u5e94\u6027\u514d\u75ab\u57fa\u56e0\u5728\u8fdb\u5316\u8fc7\u7a0b\u4e2d\u88ab\u8feb\u53d8\u6210\u4e8c\u500d\u4f53(\u5c24\u5176\u662fMHC\u2160\u7c7b\u548cMHC\u2161\u7c7b\u57fa\u56e0)\uff0c\u8fd9\u662f\u51e0\u5341\u5e74\u6765\u4e00\u4e2a\u6d3b\u8dc3\u7684\u7814\u7a76\u9886\u57df\uff0c\u5e76\u5728\u57fa\u56e0\u7ec4\u6d4b\u5e8f\u65f6\u4ee3\u5f97\u5230\u4e86\u632f\u5174\u3002<\/li>\n<li>\u6709\u4eba\u63d0\u51fa\u4e86\u4e00\u79cd\u5173\u4e8e\u9c91\u9c7c\u8fc1\u79fb\u56de\u539f\u5b75\u5316\u5730\u4ea7\u5375\u7684\u514d\u75ab\u5047\u8bf4\u3002\u5728\u7d27\u5f20\u7684\u7ec8\u672b\u8fc1\u79fb\u8fc7\u7a0b\u4e2d\uff0c\u5e7c\u7a1a\u7684\u6dcb\u5df4\u7ec6\u80de\u88ab\u8017\u5c3d\uff0c\u800c\u6d46\u7ec6\u80de\u5374\u5e78\u514d\uff0c\u5f88\u53ef\u80fd\u4ea7\u751f\u9488\u5bf9\u9c7c\u7c7b\u6700\u521d\u63a5\u89e6\u7684\u75c5\u539f\u4f53\u7684\u6297\u4f53\u3002<\/li>\n<\/ul>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p>Box 2<\/p>\n<p>Humoral immunity in the absence of germinal centres<\/p>\n<p>\u8fc7\u53bb50\u5e74\u7684\u7814\u7a76\u8868\u660e\uff0c\u603b\u7684\u6765\u8bf4\uff0c\u51b7\u8840\u810a\u690e\u52a8\u7269\u7684\u7279\u5f02\u6027\u6297\u4f53\u53cd\u5e94\u662f\u4f4e\u4eb2\u548c\u529b\u7684\uff0c\u968f\u7740\u65f6\u95f4\u7684\u63a8\u79fb\u4e0d\u4f1a\u6210\u719f\u81f3\u54fa\u4e73\u52a8\u7269\u4f53\u5185\u7684\u9ad8\u6c34\u5e73\u6297\u4f53\u3002\u5728\u5206\u5b50\u65f6\u4ee3\u4e4b\u524d\uff0c\u8fd9\u662f\u7531\u4e8e\u5728\u975e\u54fa\u4e73\u52a8\u7269\u4e2d\u53ef\u80fd\u7f3a\u4e4f\u4f53\u7ec6\u80de\u9ad8\u7a81\u53d8\u3002\u7136\u800c\uff0c\u5728\u4e8c\u5341\u4e16\u7eaa\u4e5d\u5341\u5e74\u4ee3\u65e9\u671f\uff0c\u5728\u8f6f\u9aa8\u9c7c\u7c7b\u548c\u4e24\u6816\u7c7b\u514d\u75ab\u7403\u86cb\u767d\u57fa\u56e0\u4e2d\u53d1\u73b0\u4e86\u7a81\u53d8\uff0c\u6700\u663e\u8457\u7684\u662f\u5728\u9ca8\u9c7c\u6297\u539f\u53d7\u4f53\u514d\u75ab\u7403\u86cb\u767d\u65b0\u6297\u539f\u53d7\u4f53(IgNAR)\u57fa\u56e0\u4e2d\u53d1\u73b0\u4e86\u7a81\u53d8\u3002\u5c3d\u7ba1\u5982\u6b64\uff0c\u8fc4\u4eca\u4e3a\u6b62\uff0c\u5728\u4efb\u4f55\u53d8\u6e29\u52a8\u7269\u514d\u75ab\u7814\u7a76\u4e2d\uff0c\u4eb2\u548c\u529b\u7684\u589e\u52a0\u90fd\u4e0d\u8d85\u8fc7100\u500d\uff0c\u5c3d\u7ba1\u5b83\u4eec\u81f3\u5c11\u663e\u793a\u51fa\u4e00\u5b9a\u7a0b\u5ea6\u7684\u9009\u62e9\u3002\u6b63\u5982\u672c\u6587\u6240\u63cf\u8ff0\u7684\uff0c\u81f3\u5c11\u5728\u4e24\u6816\u52a8\u7269\u7684\u5206\u7c7b\u6c34\u5e73\u4e0a\uff0c\u6ee4\u6ce1\u6811\u7a81\u72b6\u7ec6\u80de(FDC)\u5e76\u4e0d\u5b58\u5728\uff0c\u6700\u8fd1\u7684\u7814\u7a76\u8868\u660e\uff0c\u4f20\u7edf\u7684\u9020\u8840\u6e90\u6027\u6297\u539f\u63d0\u5448\u7ec6\u80de\u523a\u6fc0T\u7ec6\u80de\u548cB\u7ec6\u80de\u3002\u56e0\u6b64\uff0cFDC\u7684\u51fa\u73b0\u4e3a\u751f\u53d1\u4e2d\u5fc3\u7684\u53d1\u5c55\u63d0\u4f9b\u4e86\u6761\u4ef6\uff0c\u4e5f\u4e3a\u9ad8\u4eb2\u548c\u529b\u6210\u719f\u7684\u4ea7\u751f\u63d0\u4f9b\u4e86\u9009\u62e9\u73af\u5883\u3002\u6b63\u5982\u5148\u524d\u5173\u4e8e\u514d\u75ab\u7403\u86cb\u767d\u57fa\u56e0\u548c\u529f\u80fd\u8fdb\u5316\u7684\u7efc\u8ff0\u4e2d\u6240\u63cf\u8ff0\u7684\uff0c\u6fc0\u6d3b\u8bf1\u5bfc\u7684\u80de\u82f7\u8131\u6c28\u9176(AID)\u7684\u53d1\u73b0\u4e3a\u6dcb\u5df4\u7ec4\u7ec7\u7684\u8fdb\u5316\u63d0\u4f9b\u4e86\u4e00\u4e9b\u89c1\u89e3\uff0c\u4f46\u968f\u7740\u65f6\u95f4\u7684\u63a8\u79fb\uff0c\u5e94\u66f4\u5e7f\u6cdb\u5730\u7528\u4e8e\u7814\u7a76\u6dcb\u5df4\u7ec4\u7ec7\u4e2d\u7684\u9002\u5e94\u6027\u53cd\u5e94\u3002\u6b64\u5916\uff0c\u8fd9\u79cd\u65e0\u989a\u9c7c\u663e\u7136\u6ca1\u6709\u6b21\u7ea7\u6dcb\u5df4\u5668\u5b98\uff0c\u56e0\u6b64\u7ec6\u80de\u4e0e\u6297\u539f\u63a5\u89e6\u548c\u76f8\u4e92\u4f5c\u7528\u7684\u4f4d\u7f6e\u5c1a\u4e0d\u6e05\u695a\uff1b\u8f7d\u8102\u86cb\u767db mRNA\u7f16\u8f91\u9176\u3001\u50ac\u5316\u591a\u80bd\u6837(APOBEC)\u5bb6\u65cf\u6210\u5458\u53c2\u4e0e\u4e86\u591a\u6837\u6027\u7684\u4ea7\u751f\u548c\u53ef\u80fd\u7684\u7a81\u53d8\u3002<\/p>\n<p>Box 3<\/p>\n<p>Key questions for evolutionary immunologists<\/p>\n<p>\u6dcb\u5df4\u7ec6\u80de\u53d1\u80b2\u7684\u8f6c\u5f55\u7f51\u7edc\u662f\u5728\u9002\u5e94\u6027\u514d\u75ab\u51fa\u73b0\u4e4b\u524d\u5efa\u7acb\u7684\u3002\u8fd9\u4e2a\u7cfb\u7edf\u662f\u4f9d\u8d56\u4e8e\u5148\u5929\u6dcb\u5df4\u7ec6\u80de\u6837(ILC)\u8fc7\u7a0b\uff0c\u8fd8\u662f\u7ecf\u5178\u6dcb\u5df4\u7ec6\u80de\u9996\u5148\u51fa\u73b0\uff0c\u7136\u540e\u5931\u53bb\u6297\u539f\u53d7\u4f53\uff1f\u6ce8\u610f\uff0c\u9002\u5e94\u6027\u7ec6\u80de\u56e0\u5b50\uff0c\u5305\u62ecIL-2\u3001IL-4\u3001IFN-\u03b3\u7b49\uff0c\u5230\u76ee\u524d\u4e3a\u6b62\uff0c\u53ea\u5728\u6709\u989a\u7c7b\u4e2d\u53d1\u73b0\u3002<\/p>\n<ul>\n<li>\u542b\u6709\u514d\u75ab\u7403\u86cb\u767d\u8d85\u5bb6\u65cf(IgSF)\u548c\u4eae\u6c28\u9178\u4e30\u5bcc\u91cd\u590d\u5e8f\u5217(LRR)\u7ed3\u6784\u57df\u7684\u6297\u539f\u53d7\u4f53\u662f\u5426\u5171\u5b58\u4e8e\u4e00\u4e2a\u5171\u540c\u7684\u7956\u5148\uff1f\u662f\u4ec0\u4e48\u63a8\u52a8\u4e86\u65b0\u7cfb\u7edf\u7684\u51fa\u73b0\uff1f\u6700\u521d\u7684\u514d\u75ab\u7403\u86cb\u767d\u548cT\u7ec6\u80de\u53d7\u4f53\u529f\u80fd\u7cfb\u7edf\u6700\u521d\u662f\u57fa\u4e8e\u8f7d\u8102\u86cb\u767dB mRNA\u7f16\u8f91\u9176\u3001\u50ac\u5316\u591a\u80bd\u6837\u86cb\u767d(APOBEC)\u5bb6\u65cf\uff08\u57fa\u4e8e\u7a81\u53d8\u7684\uff09\u3001\u7136\u540e\u88ab\u57fa\u4e8e\u91cd\u7ec4\u6fc0\u6d3b\u57fa\u56e0(Rag)\u7cfb\u7edf(\u5373\u57fa\u4e8e\u91cd\u6392)\u7684\u7cfb\u7edf\u6240\u53d6\u4ee3\uff1f\u5176\u4ed6APOBEC\u5bb6\u65cf\u6210\u5458\u662f\u5982\u4f55\u5728\u6574\u4e2a\u53d8\u6e29\u52a8\u7269\u7cfb\u4e2d\u53d1\u6325\u4f5c\u7528\u7684\u5462\uff1f<\/li>\n<li>\u539fMHC\u57fa\u56e0\u7f16\u7801\u7684IgSF\u5bb6\u65cf\u6210\u5458\u662f\u5426\u4e0e\u88abRag\u8f6c\u5ea7\u5b50\u4fb5\u5165\u7684\u524d\u4f53\u57fa\u56e0\u5bc6\u5207\u76f8\u5173\uff0c\u6210\u4e3a\u539f\u59cb\u6297\u539f\u53d7\u4f53\uff1f\u5728\u7531MHC\u57fa\u56e0\u7f16\u7801\u7684\u6297\u539f\u53d7\u4f53(VJ\u578b)\u4e2d\u53d1\u73b0\u4e86\u8fd9\u79cd\u7279\u5b9a\u7c7b\u578b\u7684IgSF\u6210\u5458\u7684\u57fa\u56e0\uff0c\u5b83\u4eec\u662f\u4e0e\u8fd9\u4e00\u7956\u5148\u76f8\u5173\u7684\u826f\u597d\u5019\u9009\u57fa\u56e0\u3002<\/li>\n<li>\u4f53\u7ec6\u80de\u8d85\u7a81\u53d8\u3001\u79cd\u7c7b\u8f6c\u6362\u91cd\u7ec4\u548cT-B\u7ec6\u80de\u534f\u540c\u4f5c\u7528\u5728\u53d8\u6e29\u52a8\u7269\u751f\u53d1\u4e2d\u5fc3\u51fa\u73b0\u4e4b\u524d\u660e\u663e\u5b58\u5728\u3002\u6ee4\u6ce1\u6811\u7a81\u72b6\u7ec6\u80de\u662f\u5982\u4f55\u548c\u4e3a\u4ec0\u4e48\u5728\u9009\u62e9\u9ad8\u4eb2\u548c\u529b\u6297\u4f53\u7684\u8fc7\u7a0b\u4e2d\u88ab\u690d\u5165\u8fd9\u4e2a\u7cfb\u7edf\u7684\uff1f\u4f20\u7edf\u7684\u6811\u7a81\u72b6\u7ec6\u80de\u771f\u7684\u662f\u201c\u53cc\u91cd\u4efb\u52a1\u201d\u5411T\u548cB\u7ec6\u80de\u5448\u9012\u6297\u539f\u5417\uff1f<\/li>\n<li>\u54fa\u4e73\u52a8\u7269\u7684B1\u7ec6\u80de\u548c\u6240\u6709\u53d8\u6e29\u52a8\u7269\u5e38\u89c4B\u7ec6\u80de\u5747\u5177\u6709\u541e\u566c\u80fd\u529b\u3002\u8fd9\u662f\u5426\u610f\u5473\u7740\u9ad3\u7cfb\u548c\u6dcb\u5df4\u7cfb\u4e4b\u95f4\u5b58\u5728\u7740\u53e4\u8001\u7684\u8054\u7cfb\uff0c\u8fd9\u79cd\u673a\u5236\u5728\u51b7\u8840\u548c\u6e29\u8840\u810a\u690e\u52a8\u7269\u7684\u751f\u7406\u4e0a\u6709\u4ec0\u4e48\u7528\u5904\uff1f<\/li>\n<li>\u4e03\u9cc3\u9cd7T\u7ec6\u80de\u662f\u5982\u4f55\u8bc6\u522b\u6297\u539f\u7684\uff1f\u662f\u5426\u6709\u5916\u6765\u6297\u539f\u7684\u6536\u7f29\u80bd\u8bc6\u522b\uff0c\u8fd8\u662f\u6709\u4e00\u4e2a\u5b8c\u5168\u4e0d\u540c\u7684\u673a\u5236\uff1f<\/li>\n<li>\u4e3a\u4ec0\u4e48\u80f8\u817a\u4f1a\u8fdb\u5316\uff1f\u5982\u679c\u65e0\u989a\u7c7b\u80f8\u817a\u6837\u4f53\u662f\u4e00\u79cd\u6a21\u578b\uff0c\u4e5f\u8bb8\u6709\u989a\u7c7b\u80f8\u817a\u7684\u51fa\u73b0\u4e0d\u662f\u4e3a\u4e86\u80f8\u817a\u7684\u6b63\u5411\u6216\u8d1f\u5411\u9009\u62e9\uff0c\u800c\u662f\u4e3a\u4e86\u4f7f\u53d1\u80b2\u4e2d\u7684T\u7ec6\u80de\u8fdc\u79bbB\u7ec6\u80de\u53d1\u80b2\u7684\u5fae\u73af\u5883(\u5c31\u50cf\u9aa8\u9ad3)\u3002\u5b83(\u53ef\u80fd)\u610f\u5473\u7740\uff0c\u4e00\u65e6\u83b7\u5f97\u4e86T\u7ec6\u80de\u53d1\u80b2\u7684\u9694\u79bb\u73af\u5883\uff0c\u5c31\u4f1a\u51fa\u73b0\u590d\u6742\u7684\u6b63\u8d1f\u9009\u62e9\u673a\u5236\uff0c\u8fd9\u53ef\u80fd\u4e0eT\u7ec6\u80de\u5bf9\u80bd-MHC\u590d\u5408\u7269\u7684\u8bc6\u522b\u6709\u5173\u3002<\/li>\n<li>\u5728\u7ffb\u8bd1\u65b9\u9762\uff0c\u53ef\u53d8\u6dcb\u5df4\u7ec6\u80de\u53d7\u4f53\u548c\u9ca8\u9c7c\u5355\u6297\u662f\u8bca\u65ad\u548c\/\u6216\u6cbb\u7597\u6027\u6297\u4f53\u7684\u65b0\u5e73\u53f0\u3002\u65e0\u989a\u7c7b\u548c\/\u6216\u8f6f\u9aa8\u9c7c\u7c7b\u4e0e\u4eba\u7c7b\u4e4b\u95f4\u7684\u5de8\u5927\u7cfb\u7edf\u53d1\u80b2\u8ddd\u79bb\u5141\u8bb8\u5bf9\u4eba\u7c7b\u9776\u6807\u4e0a\u8fdb\u5316\u4fdd\u5b88\u7684\u8868\u4f4d\u4ea7\u751f\u514d\u75ab\u53cd\u5e94\u3002\u8fd9\u4e9b\u53cd\u5e94\u7269\u5728\u6211\u4eec\u7684\u836f\u5e93\u4e2d\u8bc1\u660e\u6709\u591a\u5927\u7528\u5904\uff1f<\/li>\n<li>\u4ece\u6280\u672f\u4e0a\u8bb2\uff0c\u7528\u4e8e\u57fa\u56e0\u6572\u9664\u548c\u7a81\u53d8\u7814\u7a76\u7684CRISPR\u6280\u672f\u7684\u51fa\u73b0\u5c06\u4f7f\u53d8\u6e29\u6a21\u5f0f\u751f\u7269\u7684\u57fa\u672c\u514d\u75ab\u5b66\u53d6\u5f97\u8fc5\u901f\u8fdb\u5c55\u3002\u6b64\u5916\uff0c\u65b0\u4e00\u4ee3\u5168\u57fa\u56e0\u7ec4\u548c\u8f6c\u5f55\u7ec4\u7684\u6d4b\u5e8f\u4ee5\u53ca\u86cb\u767d\u8d28\u7ec4\u7684\u5feb\u901f\u53d1\u5c55\uff0c\u81f3\u5c11\u5728\u67d0\u4e9b\u65b9\u9762\u53ef\u4ee5\u5728\u8bb8\u591a\u7814\u7a76\u4e2d\u907f\u514d\u5bf9\u6a21\u578b\u751f\u7269\u4f53\u7684\u8981\u6c42\u3002<\/li>\n<li>\u5bf9\u975e\u54fa\u4e73\u810a\u690e\u52a8\u7269\u7684\u7814\u7a76\u662f\u5426\u4f1a\u63d0\u9192\u6574\u4e2a\u514d\u75ab\u5b66\u9886\u57df\u03b3\u03b4 T\u7ec6\u80de\u7684\u9002\u5e94\u6027\u514d\u75ab\u6f5c\u529b\uff1f\u6211\u4eec\u5982\u4f55\u8bbe\u8ba1\u5b9e\u9a8c\u6765\u4e86\u89e3\u8fd9\u4e9b\u7ec6\u80de\u5982\u4f55\u8bc6\u522b\u5728\u6240\u6709\u810a\u690e\u52a8\u7269\u4e2d\u7684\u5929\u7136\u6297\u539f\uff1f<\/li>\n<li>\u5c0f\u9f20\u548c\u4eba\u7c7b\u7684\u9ecf\u819c\u514d\u75ab\u53ef\u80fd\u662f\u5b8c\u5168\u4e0d\u540c\u7684\u3002\u53d8\u6e29\u52a8\u7269\u7684\u7814\u7a76\u5982\u4f55\u8fdb\u4e00\u6b65\u4e86\u89e3\u7c98\u819c\u514d\u75ab\u7cfb\u7edf\u4e2d\u6700\u57fa\u672c\u3001\u6700\u4fdd\u5b88\u7684\u6210\u5206\uff1f<\/li>\n<li>\u8bb8\u591a\u5e74\u524d\uff0c\u6291\u5236\u7ec6\u80de\u6697\u793a\u51fa\u73b0\u5728\u4e24\u6816\u52a8\u7269\u8715\u53d8\u671f\uff0c\u4ee5\u6291\u5236\u5bf9\u6210\u866b\u7279\u5f02\u6027\u6297\u539f\u7684\u4efb\u4f55\u53cd\u5e94\u3002\u73b0\u5728\u7684\u6280\u672f\u53ef\u4ee5\u7528\u8bb8\u591a\u65b0\u7684\u8d44\u6e90\u91cd\u65b0\u5ba1\u89c6\u8fd9\u4e00\u63d0\u8bae\u3002<\/li>\n<li>\u5916\u5468\u6765\u6e90\u7684\u8c03\u8282\u6027T(Treg)\u7ec6\u80de\u771f\u7684\u662f\u4f5c\u4e3a\u8c03\u8282\u80ce\u76d8\u54fa\u4e73\u52a8\u7269\u7236\u7cfb\u7279\u5f02\u6027\u514d\u75ab\u7684\u7ec6\u80de\u4e9a\u7fa4\u51fa\u73b0\u7684\u5417\uff1f\u4e5f\u5c31\u662f\u8bf4\uff0c\u53d8\u6e29\u810a\u690e\u52a8\u7269(\u548c\u9e1f\u7c7b)\u662f\u5426\u53ea\u6709\u80f8\u817a\u6765\u6e90\u7684Treg\u7ec6\u80de\uff1f<\/li>\n<li>\u52a8\u7269\u7684\u751f\u6d3b\u65b9\u5f0f\u5728\u68c0\u67e5\u9002\u5e94\u6027\u514d\u75ab\u65f6\u663e\u7136\u5f88\u91cd\u8981\u3002\u9664\u4e86\u9cd5\u9c7c\u548c\u5357\u6781\u9c7c\u5916\uff0c\u8fd8\u6709\u8bb8\u591a\u5176\u4ed6\u79cd\u7c7b\u7684\u786c\u9aa8\u9c7c\u548c\u8f6f\u9aa8\u9c7c\u751f\u6d3b\u5728\u4e0d\u540c\u7684\u73af\u5883\u4e2d\uff0c\u5e94\u8be5\u52a0\u4ee5\u7814\u7a76\u3002<\/li>\n<li>\u54ea\u4e00\u7c7b\u5148\u51fa\u73b0\uff1aMHC\u2160\u7c7b\u8fd8\u662fMHC\u2161\u7c7b\uff1f\u5bf9\u4e8eMHC\u2160\u7c7b\uff0c\u7ecf\u5178\u5206\u5b50\u8fd8\u662f\u975e\u7ecf\u5178\u5206\u5b50\u5148\u51fa\u73b0\uff1fMHC\u2160\u7c7b\u5206\u5b50\u7684\u53ef\u5851\u6027\u8868\u660e\u5b83\u4eec\u662f\u539f\u59cb\u5206\u5b50\uff0c\u800cMHC\u2161\u7c7b\u5206\u5b50\u7684\u70ed\u529b\u5b66\u7a33\u5b9a\u6027\u662f\u5b83\u4eec\u65e9\u51fa\u73b0\u7684\u8bc1\u636e\u3002\u7c7b\u4f3c\u5730\uff0cCD4\u548cCD8\u4e0d\u662f\u6765\u81ea\u6700\u8fd1\u7684\u5171\u540c\u7956\u5148\uff0c\u6240\u4ee5\u5b83\u4eec\u8bc6\u522bMHC\u5206\u5b50\u7684\u5171\u540c\u9009\u62e9\u662f\u72ec\u7acb\u7684\u3002\u54ea\u4e2a\u662f\u7b2c\u4e00\u4f4d\u7684\uff1f<\/li>\n<\/ul>\n<p>&nbsp;<\/p>\n<p>\u7248\u6743\u58f0\u660e\uff1a\u672c\u6587\u4e3a\u535a\u4e3b\u539f\u521b\u6587\u7ae0\uff0c\u672a\u7ecf\u535a\u4e3b\u5141\u8bb8\u4e0d\u5f97\u8f6c\u8f7d\u3002<\/p>\n<p>\u539f\u6587\uff1a<a href=\"https:\/\/www.nature.com\/articles\/s41577-018-0003-9\">https:\/\/www.nature.com\/articles\/s41577-018-0003-9<\/a><\/p>\n","protected":false},"excerpt":{"rendered":"<p>Abstract\uff1a\u9002\u5e94\u6027\u514d\u75ab\u7cfb\u7edf\u8d77\u6e90\u4e8e5\u4ebf\u5e74\u524d\u7684\u53d8\u6e29(\u51b7\u8840)\u810a\u690e\u52a8\u7269\u3002\u4f20\u7edf\u4e0a\uff0c\u9002\u5e94\u6027\u514d\u75ab\u7cfb\u7edf\u662f\u7531\u8868\u8fbe\u91cd\u7ec4\u6fc0\u6d3b\u57fa\u56e0(Rag)\u4f9d\u8d56\u6297\u539f\u53d7\u4f53\u7684\u6dcb\u5df4\u7ec6\u80de\u548cMHC\u7ec4\u6210\u7684\u3002\u8fd9\u4e9b\u7279\u5f81\u5b58\u5728\u4e8e\u6240\u6709\u4e0b\u988c\u9aa8\u7684\u810a\u690e\u52a8\u7269\u4e2d\uff0c\u5305\u62ec\u8f6f\u9aa8\u548c\u786c\u9aa8\u9c7c\u3001\u4e24\u6816\u52a8\u7269\u548c\u722c\u884c\u52a8\u7269\uff0c\u53ef\u80fd\u4e5f\u5b58\u5728\u4e8e\u6700\u53e4\u8001\u7684\u4e0b\u989a\u810a\u690e\u52a8\u7269\uff0c\u5373\u706d\u7edd\u7684\u80ce\u76d8\u76ae\u52a8\u7269\u4e2d\u3002\u7136\u800c\uff0c\u968f\u7740\u65e0\u988c\u9c7c\u7c7b\u9002\u5e94\u6027\u514d\u75ab\u7cfb\u7edf\u7684\u53d1\u73b0\uff0c\u57fa\u4e8e\u4e00\u7ec4\u5b8c\u5168\u4e0d\u540c\u7684\u6297\u539f\u53d7\u4f53-\u53ef\u53d8\u6dcb\u5df4\u7ec6\u80de\u53d7\u4f53-t\u548cb\u7ec6\u80de\u7684\u5206\u53c9\uff0c\u4e5f\u8bb8\u662f\u5148\u5929\u6837\u6dcb\u5df4\u7ec6\u80de\uff0c\u53ef\u4ee5\u8ffd\u6eaf\u5230\u6240\u6709\u810a\u690e\u52a8\u7269\u7684\u8d77\u6e90\u3002\u672c\u7efc\u8ff0\u63a2\u8ba8\u6bd4\u8f83\u514d\u75ab\u5b66\u7684\u6700\u65b0\u8fdb\u5c55\uff0c\u4fc3\u8fdb\u6211\u4eec\u5bf9\u9002\u5e94\u6027\u514d\u75ab\u7cfb\u7edf\u7684\u8d77\u6e90\u548c\u529f\u80fd\u7684\u7406\u89e3\u3002 Introduction\uff1a\u5927\u591a\u6570\u5b9a\u4e49\u4e86\u514d\u75ab\u5b66\u9886\u57df\u7684\u7814\u7a76\u90fd\u662f\u5728\u54fa\u4e73\u52a8\u7269\u8eab\u4e0a\u8fdb\u884c\u7684\uff0c\u7279\u522b\u662f\u5728\u8001\u9f20\u548c\u4eba\u7c7b\u8eab\u4e0a\uff0c\u800c\u6211\u4eec\u7684\u5927\u90e8\u5206\u89c4\u5f8b\u548c\u8303\u5f0f\u90fd\u662f\u4ece\u8fd9\u4e9b\u6a21\u578b\u4e2d\u884d\u751f\u51fa\u6765\u7684\u3002\u7136\u800c\uff0cJaneway and Matzinger\u5236\u5b9a\u4e86\u4e00\u6761\u65b0\u7684\u6cd5\u5219\uff0c\u5b83\u8bf4\u660e\u9002\u5e94\u6027\u514d\u75ab\u7684\u884c\u4e3a\u4e0d\u662f\u7531\u5916\u6765\uff0c\u800c\u662f\u901a\u8fc7\u6a21\u5f0f\u8bc6\u522b\u53d7\u4f53(PRRs)\u88ab\u5916\u90e8\u6216\u5185\u90e8\u5371\u9669\u6240\u6fc0\u53d1\u7684\uff0c\u5b83\u8d77\u6e90\u4e8e\u679c\u8747\u9ed1\u8179\u9f20Toll-like receptors\u7684\u7814\u7a76\u3002\u6b64\u5916\uff0c50\u591a\u5e74\u524d\uff0c\u5728\u9e1f\u7c7b\u6cd5\u6c0f\u56ca\u7684\u7814\u7a76\u4e2d\u53d1\u73b0\u4e86\u4e24\u4e2a\u4e3b\u8981\u6dcb\u5df4\u7ec6\u80de\u4e9a\u7fa4(B\u7ec6\u80de\u548cT\u7ec6\u80de)\u7684\u5dee\u5f02\u3002\u4e3a\u4e86\u4e86\u89e3\u9002\u5e94\u6027\u514d\u75ab\u7684\u8d77\u6e90\uff0c\u6211\u4eec\u5fc5\u987b\u4e86\u89e3\u53d8\u6e29\u810a\u690e\u52a8\u7269\uff0c\u4ee5\u53ca\u810a\u690e\u52a8\u7269\u7684\u76f4\u63a5\u7956\u5148\u2014\u8f83\u4f4e\u7b49\u7684\u540e\u53e3\u52a8\u7269\u3002\u5728\u8fc7\u53bb\u7684\u5341\u5e74\u91cc\uff0c\u7a81\u7834\u6027\u7684\u53d1\u73b0\u5e26\u6765\u4e86\u4eba\u4eec\u5bf9\u9002\u5e94\u6027\u514d\u75ab\u8d77\u6e90\u7684\u8d8a\u6765\u8d8a\u591a\u7684\u8ba4\u8bc6\uff0c\u5e76\u63d0\u9192\u6211\u4eec\u6ce8\u610f\u65b0\u7684\u53ef\u80fd\u6027\u3002 \u4e86\u89e3\u514d\u75ab\u8fdb\u5316\u7684\u4e00\u4e2a\u65b9\u4fbf\u7684\u65b9\u6cd5\u662f\u5c06\u514d\u75ab\u7cfb\u7edf\u5212\u5206\u4e3a\u968f\u7740\u65f6\u95f4\u7684\u63a8\u79fb\u800c\u4fdd\u5b88\u7684\u90e8\u5206\uff0c\u800c\u4e0d\u662f\u90a3\u4e9b\u53d8\u5316\u8fc5\u901f\u7684\u90e8\u5206\uff0c\u8fd9\u662fJan Klein\u6700\u521d\u63d0\u51fa\u7684\u60f3\u6cd5\u3002\u67d0\u4e9b\u514d\u75ab\u7279\u5f81\uff0c\u5982\u514d\u75ab\u7403\u86cb\u767dM(IgM)\u7684\u7ed3\u6784\u548c\u529f\u80fd\uff0c\u4ee5\u53ca\u80f8\u817a\u3001\u813e\u810f\u3001\u5e38\u89c4\u03b1\u03b2T\u7ec6\u80de\u53d7\u4f53(TCRs)\u548cMHC\u2161\u7c7b\u5206\u5b50\u7684\u5b58\u5728\uff0c\u5c06\u5728\u672c\u7efc\u8ff0\u4e2d\u8f83\u4e3a\u8be6\u7ec6\u5730\u63cf\u8ff0\uff0c\u5728\u51e0\u4e4e\u6240\u6709\u7684\u6709\u989a\u7c7b\u810a\u690e\u52a8\u7269\uff08gnathostomes\uff09\u4e2d\u90fd\u662f\u9ad8\u5ea6\u4fdd\u5b88\u7684\u3002\u76f8\u53cd\uff0c\u5176\u4ed6\u514d\u75ab\u6210\u5206\uff0c\u5982IgD\u3001\u03b3\u03b4TCR\u3001\u81ea\u7136\u6740\u4f24\u53d7\u4f53(NKRs)\u548c\u975e\u7ecf\u5178\u7684MHC\u5206\u5b50\u90fd\u662f\u53ef\u5851\u6027\u7684\uff0c\u5728\u57fa\u56e0\u6570\u76ee\uff0c\u7ed3\u6784\u57df\u548c\u529f\u80fd\u4e0a\u603b\u662f\u5b58\u5728\u5dee\u5f02\u3002\u8bb0\u4f4f\u8fd9\u4e00\u6807\u51c6\u4e3a\u7406\u89e3\u514d\u75ab\u7cfb\u7edf\u7684\u57fa\u7840\u63d0\u4f9b\u4e86\u4e00\u4e2a\u6846\u67b6\uff1a\u4fdd\u7559\u7ecf\u8fc7\u9a8c\u8bc1\u7684\u4e8b\u5b9e\uff0c\u4f46\u5141\u8bb8\u8fdb\u5316\u4e0a\u7684\u5feb\u901f\u53d8\u5316\u4e0e\u5176\u4ed6\u4e92\u8865\u7279\u5f81\uff0c\u4ee5\u5bf9\u6297\u4e0d\u65ad\u53d8\u5316\u7684\u75c5\u539f\u4f53\uff08\u73af\u5883\uff09\u3002 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2\uff09\u3002 \u65e0\u988c\u9c7c\u6709T\u7ec6\u80de(both \u03b1\u03b2 and \u03b3\u03b4 T cells) \u548cB\u7ec6\u80de\uff0c\u4ee5\u53ca\u80f8\u817a\u7c7b\u4f3c\u7269\uff0cMHC\u88ab\u8bc1\u660e\u662f\u96be\u4ee5\u627e\u5230\u7684\uff08Fig 1\u30012\uff09\u3002\u5c31\u50cf\u53ef\u53d8\u6dcb\u5df4\u7ec6\u80de\u53d7\u4f53(VLRs)\uff0cMHC\u53ef\u80fd\u662f\u901a\u8fc7\u6536\u655b\u6f14\u5316\u5728\u8fd9\u4e2a\u7fa4\u4f53\u4e2d\u4ea7\u751f\u7684\u3002\u6bcf\u4e00\u7ec4\u4e2d\u7c98\u819c\u9002\u5e94\u6027\u514d\u75ab\u662f\u5b58\u5728\u7684\u548c\u72ec\u7279\u7684\uff0c\u5982\u4e24\u6816\u52a8\u7269\u548c\u786c\u9aa8\u9c7c\u7684\u7814\u7a76(\u548c\u54fa\u4e73\u52a8\u7269)\u3002\u5728\u4e9a\u5206\u7c7b\u7fa4\u4e2d\u53d1\u73b0\u8f6f\u9aa8\u9c7c\u3001\u786c\u9aa8\u9c7c\u548c\u4e24\u6816\u52a8\u7269\u663e\u793a\u51fa\u533a\u522b\u6574\u4e2a\u810a\u690e\u52a8\u7269\u95e8\u7684\u7279\u5f81\u4f46\u4e5f(\u5728\u67d0\u4e9b\u60c5\u51b5\u4e0b)\u6709\u7740\u72ec\u7279\u7684\u7279\u5f81\uff08Fig 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\u2161\u7c7b\u7cfb\u7edf\u7684\u7f3a\u5931\u548c\u67d0\u4e9b\u514d\u75ab\u7403\u86cb\u767d\u4e9a\u578b\u7684\u7f3a\u5931\u3002\u8f6f\u9aa8\u9c7c\u5177\u6709\u72ec\u7279\u7684\u514d\u75ab\u7403\u86cb\u767d\u7ec4\u7ec7\uff0c\u4f7f\u5f97\u6297\u539f\u53d7\u4f53\u57fa\u56e0\u7684\u51fa\u73b0\u5177\u6709\u65b0\u7684\u529f\u80fd\u3002\u867d\u7136NK\u7ec6\u80de\u660e\u663e\u5b58\u5728\u4e8e\u53d8\u6e29\u52a8\u7269\uff0c\u7531\u4e8e\u514d\u75ab\u7cfb\u7edf\u7684\u5feb\u901f\u53d1\u5c55\uff0c\u8bc6\u522bNKRs\u53d8\u5f97\u975e\u5e38\u56f0\u96be\u3002\u7136\u800c\uff0c\u4e0eMHC\u76f8\u5173\u7684NKR\u57fa\u56e0\u5df2\u5728\u6240\u6709\u810a\u690e\u52a8\u7269\u4e2d\u53d1\u73b0\uff0c\u663e\u793a\u51faMHC \u2160\u548cNKRs\u65e9\u671f\u76f8\u5173\u6027\u3002\u5728\u8fc4\u4eca\u7814\u7a76\u7684\u6240\u6709\u9c7c\u7c7b\u548c\u4e24\u6816\u52a8\u7269\u7684MHC\u4e2d\u90fd\u53d1\u73b0\u4e86\u7f16\u7801MHC\u2160\u7c7b\u5206\u5b50\u3001\u514d\u75ab\u86cb\u767d\u9176\u4f53\u548c\u4e0e\u6297\u539f\u52a0\u5de5\u5206\u5b50\u76f8\u5173\u7684\u8f6c\u8fd0\u4f53(TAP)\u57fa\u56e0\u7684\u53e4\u8001\u8c31\u7cfb\uff0c\u4f46\u662f\u5728\u54fa\u4e73\u52a8\u7269\u4e2d\uff0c\u8fd9\u4e2a\u539f\u578b\u5df2\u7ecf\u88ab\u4e00\u4e2a\u5728MHC\u2160\u7c7b\u80bd\u7279\u5f02\u6027\u4e0d\u90a3\u4e48\u4e25\u683c\u7684\u7cfb\u7edf\u6240\u53d6\u4ee3\u3002\u8fdb\u5316\u4e0a\u7684\u4e00\u5927\u98de\u8dc3\u5728\u4e24\u6816\u52a8\u7269\u4e2d\u662f\u660e\u663e\u7684\uff0c\u5b83\u4eec\u8868\u73b0\u51fa\u5178\u578b\u7684\u6297\u4f53\u7c7b\u8f6c\u6362\uff0c\u5e76\u4e14\u6709\u4e00\u4e2aIgG\u7c7b(\u79f0\u4e3aIgY)\uff0c\u5b83\u53c2\u4e0e\u5178\u578b\u7684\u8bb0\u5fc6\u514d\u75ab\u53cd\u5e94\uff08Fig 2\uff09\u3002\u810a\u690e\u52a8\u7269\u8fdb\u5316\u7684\u6700\u540e\u4e00\u4e2a\u91cd\u5927\u8fdb\u5c55\u662f\u6dcb\u5df4\u7ed3\u7684\u51fa\u73b0\u548c\u54fa\u4e73\u52a8\u7269\u751f\u53d1\u4e2d\u5fc3\u7684\u5f62\u6210\uff1b\u53d8\u6e29\u810a\u690e\u52a8\u7269\u5141\u8bb8\u5728\u6ee4\u6ce1\u6811\u7a81\u72b6\u7ec6\u80de(FDCs)\u51fa\u73b0\u4e4b\u524d\u8fdb\u884c\u514d\u75ab\u7814\u7a76\uff0c\u800c\u6811\u7a81\u72b6\u7ec6\u80de\u662f\u54fa\u4e73\u52a8\u7269affinity\u6210\u719f\u7684\u4e3b\u8981\u7ec6\u80de\u3002\u672c\u7efc\u8ff0\u5c06\u6df1\u5165\u7814\u7a76\u9002\u5e94\u6027\u514d\u75ab\u7684\u6240\u6709\u8fd9\u4e9b\u7279\u6027\uff0c\u91cd\u70b9\u8ba8\u8bba\u5f15\u8d77\u6240\u6709\u514d\u75ab\u5b66\u5bb6\u5174\u8da3\u7684\u57fa\u672c\u95ee\u9898\uff0c\u5e76\u63a2\u8ba8\u6211\u4eec\u5728\u672a\u6765\u5341\u5e74\u4e2d\u80fd\u591f\u548c\u5e94\u8be5\u89e3\u51b3\u7684\u95ee\u9898\u3002\u503c\u5f97\u6ce8\u610f\u7684\u662f\uff0c\u6e29\u8840\u6027\u662f\u5728\u4f4e\u7b49\u810a\u690e\u52a8\u7269\u7684\u51e0\u4e2a\u5206\u7c7b\u7fa4\u4e2d\u53d1\u73b0\u7684\uff0c\u4f46\u603b\u7684\u6765\u8bf4\uff0c\u5b83\u4eec\u7684\u9002\u5e94\u6027\u514d\u75ab\u8fd8\u6ca1\u6709\u88ab\u68c0\u6d4b\u3002\u5728\u8fd9\u91cc\uff0c\u6211\u5c06\u96c6\u4e2d\u4e8e\u9c7c\u7c7b\u3001\u4e24\u6816\u7c7b\u548c\u722c\u884c\u52a8\u7269\u65b9\u9762\u7684\u5927\u90e8\u5206\u7814\u7a76\u3002 Fig 1. Fig 2. Evolution of antigen receptors \u989a\u53e3\u7c7b\u7684\u6297\u4f53\u548cTCRs\u662f\u514d\u75ab\u7403\u86cb\u767d\u8d85\u5bb6\u65cf\uff08IgSF\uff09\u7684\u6210\u5458\uff0c\u6765\u6e90\u4e8e\u672a\u77e5\u7684\u524d\u4f53\u3002\u63d0\u51fa\u4e86\u4e00\u4e2a\u901a\u7528\u6a21\u578b\uff0c\u5e76\u5728\u4e0b\u9762\u8ba8\u8bba\u3002\u76f8\u6bd4\u4e4b\u4e0b\uff0c\u65e0\u988c\u9c7cVLRs\u5c5e\u4e8e\u53e4\u8001\u7684\u5bcc\u542b\u4eae\u6c28\u9178\u7684\u91cd\u590d\u5e8f\u5217(LRR)\u53d7\u4f53\u5bb6\u65cf\uff0c\u53ef\u80fd\u8d77\u6e90\u4e8e\uff08\u4e5f\u662f\u6700\u76f8\u5173\u7684\u4e00\u79cd\u53d7\u4f53\uff09\u8840\u5c0f\u677f\u4e0a\u8868\u8fbe\u7684\u4e00\u79cd\u7ec6\u80de\u8868\u9762\u53d7\u4f53\u3002\u5728\u6dcb\u5df4\u7ec6\u80de\u53d1\u80b2\u8fc7\u7a0b\u4e2d\uff0c\u4e24\u79cd\u6297\u539f\u53d7\u4f53\u90fd\u53ef\u4ee5\u4ea7\u751f\u5f88\u9ad8\u6c34\u5e73\u7684\u591a\u6837\u6027\uff0c\u4f46\u7528\u4e8e\u4ea7\u751f\u591a\u6837\u6027\u7684\u57fa\u56e0\u91cd\u6392\u673a\u5236\u5219\u5b8c\u5168\u4e0d\u540c\u3002 Variable lymphocyte receptors \u4e4b\u524d\u5bf9\u4f53\u6db2\u514d\u75ab\u548c\u7ec6\u80de\u514d\u75ab\u8fdb\u884c\u7684\u4e00\u4e9b\u7814\u7a76\u8868\u660e\uff0c\u65e0\u989a\u7c7b\u5177\u6709\u9002\u5e94\u6027\u514d\u75ab\uff0c\u5bf9\u4e0d\u540c\u7684\u5916\u6765\u6297\u539f\u548c\u540c\u79cd\u6297\u539f\u6709\u7279\u5f02\u6027\u7684\u5e94\u7b54\u3002\u7136\u800c\uff0c30\u591a\u5e74\u6765\u5bf9\u6297\u4f53\u3001TCRs\u548cMHC\u5206\u5b50\u7684\u5e7f\u6cdb\u7814\u7a76\u88ab\u8bc1\u660e\u662f\u5f92\u52b3\u7684\uff0c\u5927\u591a\u6570\u6bd4\u8f83\u514d\u75ab\u5b66\u5bb6\u90fd\u8ba4\u4e3a\uff0c\u5c3d\u7ba1\u65e9\u671f\u8fdb\u884c\u4e86\u529f\u80fd\u7814\u7a76\uff0c\u4f46\u65e0\u989a\u7c7b\u6ca1\u6709\u9002\u5e94\u6027\u514d\u75ab\u7cfb\u7edf\u5b58\u5728\u3002\u8fd9\u4e00\u5047\u8bbe\u88abPancer\u548cCooper\u8bc1\u660e\u662f\u4e0d\u6b63\u786e\u7684\uff0c\u4ed6\u4eec\u68c0\u67e5\u4e86\u514d\u75ab\u4e86\u7684\u4e03\u9cc3\u9cd7\u4e2d\u6dcb\u5df4\u7ec6\u80de\u7684\u8f6c\u5f55\u7ec4\uff0c\u5e76\u53d1\u73b0\u4e86\u5927\u91cf\u542b\u6709\u4e0d\u540c\u6570\u91cf\u5185\u90e8\u91cd\u590d\u5e8f\u5217\u7684LRR\u86cb\u767d\uff08Fig 3\uff09\u3002\u8fd9\u4e9b\u542bLRR\u7684\u53d7\u4f53\u5728\u6dcb\u5df4\u7ec6\u80de\u4e2a\u4f53\u53d1\u751f\u8fc7\u7a0b\u4e2d\u88ab\u514b\u9686\u8868\u8fbe\u5e76\u901a\u8fc7\u91cd\u6392\u4ea7\u751f\uff0c\u5e76\u88ab\u547d\u540d\u4e3aVLRs\u3002\u7b2c\u4e00\u4e2a\u8981\u7814\u7a76\u7684\u57fa\u56e0\u662fVLRB\uff0c\u8be5\u57fa\u56e0\u5728\u4e00\u7ec4\u4e03\u9cc3\u9cd7\u6dcb\u5df4\u7ec6\u80de\u4e2d\u8868\u8fbe\u3002VLRB\u86cb\u767d\u88ab\u7cd6\u57fa\u78f7\u8102\u9170\u808c\u9187\u951a\u5b9a\u5728\u4e03\u9cc3\u9cd7\u521a\u5f62\u6210\u7684\u6dcb\u5df4\u7ec6\u80de\u8868\u9762\uff0c\u7136\u540e(\u901a\u8fc7\u672a\u77e5\u7684\u673a\u5236)\u5728\u6297\u539f\u523a\u6fc0\u540e\u4ee5\u4e8c\u805a\u4f53\u7684\u4e94\u805a\u4f53\u5f62\u5f0f\u5206\u6ccc\uff08Fig 3\uff09\u3002\u9996\u5148\uff0c\u4eba\u4eec\u8ba4\u4e3a\u523a\u6fc0\u4e03\u9cc3\u9cd7\u6dcb\u5df4\u7ec6\u80de\u7c7b\u4f3c\u4e8e\u54fa\u4e73\u52a8\u7269\u7684\u975eT\u7ec6\u80de\u4f9d\u8d56\u7684\u514d\u75ab\u53cd\u5e94\uff0c\u5373\u6dcb\u5df4\u7ec6\u80de\u4e0a\u6297\u539f\u53d7\u4f53\u7684\u4ea4\u8054\uff0c\u5f88\u53ef\u80fd\u901a\u8fc7PRR\u4e0e\u53e6\u4e00\u4e2a\u4fe1\u53f7\u7ed3\u5408\u5728\u4e00\u8d77\uff0c\u53ef\u80fd\u6fc0\u6d3bB\u7ec6\u80de\u4ee5\u8bf1\u5bfcVLRB\u5206\u6ccc\u3002\u7136\u800c\uff0c\u4ee4\u4eba\u60ca\u8bb6\u7684\u662f\uff0c\u540e\u6765\u53d1\u73b0VLR\u57fa\u56e0\u4f4d\u70b9\u7b2c\u4e8c\u6b21\u91cd\u6392\uff0cVLRA\u57fa\u56e0\u4f4d\u70b9\u7531\u53e6\u4e00\u90e8\u5206\u6dcb\u5df4\u7ec6\u80de\u8868\u8fbe\uff0c\u5176\u8f6c\u5f55\u7ec4\u4e0e\u6709\u989a\u7c7bT\u7ec6\u80de\u76f8\u4f3c\u3002\u8fdb\u4e00\u6b65\u5206\u6790\u53d1\u73b0\uff0c\u7b2c\u4e09\u79cd\u6297\u539f\u53d7\u4f53VLRC\u4e3b\u8981\u5728\u4e0a\u76ae\u548c\u7c98\u819c\u4e2d\u8868\u8fbe\uff0c\u56e0\u6b64\u65e0\u989a\u7c7bT\u7ec6\u80de\u4f3c\u4e4e\u88ab\u5206\u88c2\u6210\u4e24\u4e2a\u4e9a\u7fa4\uff0c\u53ef\u80fd\u7c7b\u4f3c\u6709\u989a\u7c7b\u03b3\u03b4 \u548c \u03b1\u03b2\u7ec6\u80de\u3002 Fig 3. IgM antibodies \u5728\u6240\u6709\u7684\u6709\u989a\u7c7b\u4e2d\u53d1\u73b0\u591a\u79cd\u6297\u4f53\u4e9a\u578b\u3002\u591a\u5e74\u6765\uff0c\u5c3d\u7ba1\u5728\u6bcf\u4e2a\u810a\u690e\u52a8\u7269\u7c7b\u4e2dIgM\u6709\u5176\u72ec\u7279\u800c\u8010\u4eba\u5bfb\u5473\u7684\u7279\u5f81\uff0cIgM\u4e00\u76f4\u88ab\u8ba4\u4e3a\u662f\u539f\u59cb\u6297\u4f53\u7c7b\u3002\u5728\u54fa\u4e73\u52a8\u7269\u4e2d\uff0cIgM\u662f\u4e00\u79cd\u4e0e\u8fde\u63a5(J)\u94fe\u76f8\u5173\u7684\u4e94\u805a\u4f53\u3002\u8f6f\u9aa8\u9c7c\u4e2d\u7684IgM\u6709\u4e24\u79cd\u5f62\u5f0f\uff1a\u5c31\u50cf\u6240\u6709\u5176\u4ed6\u810a\u690e\u52a8\u7269\u4e00\u6837\uff0c\u5b58\u5728\u7684\u4e00\u79cd\u591a\u91cd\u5f62\u5f0f\uff0c\u4ee5\u53ca\u4e00\u79cd\u5355\u4f53\u5f62\u5f0f\uff0c\u5728\u9ca8\u9c7c\u6297\u539f\u7279\u5f02\u6027\u53cd\u5e94\u4e2d\u6700\u5e38\u89c1\uff0c\u5f88\u53ef\u80fd\u662f\u4ee5T\u7ec6\u80de\u4f9d\u8d56\u7684\u65b9\u5f0f\u4ea7\u751f\u7684\uff08\u8bf7\u6ce8\u610f\uff0c\u867d\u7136\u5728\u6240\u6709\u5176\u4ed6\u810a\u690e\u52a8\u7269\u4e2d\u4e5f\u53d1\u73b0\u4e86\u5355\u805aIGM\uff0c\u4f46\u53ea\u6709\u5728\u8f6f\u9aa8\u9c7c\u4e2d\u624d\u53d1\u73b0\u5b83\u5728\u514d\u75ab\u5e94\u7b54\u8fc7\u7a0b\u4e2d\u5177\u6709\u91cd\u8981\u7684\u751f\u7406\u76f8\u5173\u6027\uff09\u3002\u786c\u9aa8\u9c7c\u7c7bIgM\u7684\u5206\u6ccc\u5f62\u5f0f\u662f\u4e00\u79cd\u56db\u805a\u4f53\uff0c\u800cj\u94fe\u57fa\u56e0\u5728\u8fd9\u4e00\u7ec4\u4e2d\u5df2\u7ecf\u4e22\u5931\uff0c\u56e0\u4e3a\u5b83\u5b58\u5728\u4e8e\u53e4\u8001\u7684\u8f6f\u9aa8\u9c7c\u4e2d\u3002\u6709\u8da3\u7684\u662f\uff0c\u786c\u9aa8\u9c7c\u5206\u6ccc\u7684IgM\u4e2d\u7684\u4e8c\u786b\u952e\u7684\u8303\u56f4\u662f\u5728\u6297\u539f\u7279\u5f02\u6027\u53cd\u5e94\u7684\u8fc7\u7a0b\u4e2d\u4fee\u6539\u7684\uff0c\u8fd9\u79cd\u4eb2\u548c\u529b\u7684\u6210\u719f\u4f3c\u4e4e\u662f\u8fd9\u7c7b\u6297\u4f53\u6240\u72ec\u6709\u7684\u3002\u6b64\u5916\uff0c\u786c\u9aa8\u9c7cIgM\u7684\u8de8\u819c\u5f62\u5f0f\u4e5f\u662f\u4ea4\u66ff\u62fc\u63a5\u7684\uff0c\u56e0\u6b64\u7531\u4e8e\u672a\u77e5\u7684\u529f\u80fd\u539f\u56e0\uff0c\u5b83\u53ea\u6709\u4e09\u4e2a\u4fdd\u5b88(C)\u7ed3\u6784\u57df(\u800c\u4e0d\u662f\u901a\u5e38\u7684\u56db\u4e2a)\u3002\u603b\u4e4b\uff0cIgM\u662f\u786c\u9aa8\u9c7c\u7c7b\u4e2d\u53d1\u73b0\u7684\u4e3b\u8981\u8840\u6e05\u6297\u4f53\uff0c\u662f\u8fd9\u4e9b\u52a8\u7269\u5bf9\u5e38\u89c4\u6297\u539f\u7684\u53cd\u5e94\uff1b\u8f6f\u9aa8\u9c7c\u9664\u4e86\u5728\u9002\u5e94\u6027\u53cd\u5e94\u4e2d\u4f7f\u7528IgM\u4f5c\u4e3a\u5148\u5929\u6297\u4f53\u7c7b\u5916\uff0c\u8fd8\u5728\u9002\u5e94\u6027\u514d\u75ab\u4e2d\u4f7f\u7528\u5176\u4ed6\u6297\u4f53\u4e9a\u578b\uff1b\u4e24\u6816\u7c7bIgM(\u636e\u6211\u4eec\u6240\u77e5)\u7684\u529f\u80fd\u4e0e\u5176\u54fa\u4e73\u52a8\u7269\u7684\u540c\u6e90\u57fa\u56e0\u975e\u5e38\u76f8\u4f3c\u3002 IgD and IgW antibodies IgD\uff0c\u957f\u671f\u4ee5\u6765\u88ab\u8ba4\u4e3a\u662f\u810a\u690e\u52a8\u7269\u4e2d\u65b0\u53d1\u73b0\u7684\u4e00\u79cd\uff0c\u5b9e\u9645\u4e0a\u975e\u5e38\u53e4\u8001\uff0c\u5b83\u53ef\u4ee5\u8ffd\u6eaf\u5230\u6709\u989a\u7c7b\u7684\u8d77\u6e90\uff0c\u4e00\u79cd\u5728\u8f6f\u9aa8\u9c7c\u4e2d\u53d1\u73b0\u7684\u540c\u79cd\u5f02\u578b\uff0c\u53eb\u505aIgW\uff08Fig 3\uff09\u3002\u4e00\u79cd\u7c7b\u4f3c\u4e8eIgD\u7684\u514d\u75ab\u7403\u86cb\u767d\u5728\u786c\u9aa8\u9c7c\u7c7b\u4e2d\u9996\u5148\u88ab\u53d1\u73b0\uff0c1998\u5e74\u4e4b\u524d\uff0cIgD\u53ea\u5728\u4e00\u4e9b\u54fa\u4e73\u52a8\u7269\u4e2d\u88ab\u53d1\u73b0\uff0c\u8fd9\u662f\u975e\u5e38\u4ee4\u4eba\u60ca\u8bb6\u7684\u3002\u968f\u540e\uff0c\u5728\u4e24\u6816\u52a8\u7269\u5c5e\u975e\u6d32\u722a\u87fe\uff08Xenopus\uff09\u57fa\u56e0\u7ec4\u6570\u636e\u5e93\u4e2d\u53d1\u73b0\u4e86IgD\u57fa\u56e0\uff0c\u5e76\u4e0e\u54fa\u4e73\u52a8\u7269\u4e00\u6837\uff0c\u901a\u8fc7IgM\u548cIgD mRNA\u5728\u65b0\u751fB\u7ec6\u80de\u4e2d\u7684\u9009\u62e9\u6027\u526a\u63a5\u6765\u8868\u8fbe\u3002IgD\u3001IgW\u5728\u8f6f\u9aa8\u9c7c\u7c7b\u7684\u540c\u6e90\u6027\u4e5f\u5728\u80ba\u9c7c\uff08lungfish\uff09\u4e2d\u53d1\u73b0\uff0c\u5e76\u4e14\u5728\u8154\u68d8\u9c7c\uff08coelacanths\uff09\u4e2d\uff0c\u5b83\u4f3c\u4e4e\u662f\u4e3b\u8981\u7684\u540c\u5de5\u578b\uff0c\u8fd9\u79cd\u8457\u540d\u7684\u7269\u79cd\u5df2\u7ecf\u5931\u53bb\u4e86IgM\u3002\u6700\u8fd1\u6709\u5173\u786c\u9aa8\u9c7c\u7c7b\u548c\u4eba\u7c7b\u7684\u6570\u636e\u8868\u660e\uff0cIgD\u53c2\u4e0e\u708e\u75c7\u53cd\u5e94\uff0c\u4e0e\u55dc\u78b1\u6027\u7ec6\u80de\u4e0a\u4e00\u4e2a\u672a\u77e5\u7684Fc\u53d7\u4f53\u7ed3\u5408\u3002\u4e0eIgM\u76f8\u6bd4\uff0cIgD\u5728\u7ed3\u6784\u548c\u529f\u80fd\u4e0a\u90fd\u6709\u5f88\u5927\u7684\u53ef\u5851\u6027\uff1bIgM\u5728\u6574\u4e2a\u8fdb\u5316\u8fc7\u7a0b\u4e2d\u662f\u975e\u5e38\u76f8\u4f3c\u7684\uff0c\u800cIgD\u5728C\u7ed3\u6784\u57df\u7684\u6570\u76ee\u4e0a(\u751a\u81f3\u5728IgD\u672c\u8eab\u7684\u5b58\u5728\u4e0b)\u662f\u9ad8\u5ea6\u53ef\u53d8\u7684\uff0c\u5728\u529f\u80fd\u4e0a\u53ef\u80fd\u4e0e\u8de8\u819c(\u5927\u90e8\u5206\u672a\u77e5)\u548c\u5206\u6ccc(\u56fa\u6709\u7ec6\u80de\u6b66\u88c5)\u7684\u5f62\u5f0f\u6709\u5173\u3002 IgG and IgY antibodies 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