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            <h1 id="seo-header">实用免疫学（六）适应性免疫：细胞免疫二 CTL细胞</h1>
            
            
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                <p>DC细胞提呈外源性抗原激活Th细胞，对于内源性抗原和进入细胞内的抗原，则通过所有有核细胞的MHCI类分子进行呈递。（病毒感染细胞合成病毒蛋白、肿瘤细胞合成肿瘤蛋白、吞噬体中逃逸至细胞质中的抗原、ER中折叠错误的自身蛋白）</p>
<h2 id="抗原肽-MHCI复合物的形成"><a href="#抗原肽-MHCI复合物的形成" class="headerlink" title="抗原肽-MHCI复合物的形成"></a>抗原肽-MHCI复合物的形成</h2><ul>
<li><p>第一步 内源性抗原降解成抗原肽</p>
</li>
<li><p>胞质内的靶蛋白和泛素结合，被蛋白酶体的帽子结构识别，被剪切成多肽</p>
</li>
<li><p>蛋白酶体为中空圆柱体结构，分为具有酶活性的两个β内环和不具有酶活性的两个α外环，β内环中有三个具有含有水解蛋白的活性位点亚基</p>
</li>
<li><p>在IFN-γ刺激下，蛋白酶体β内环中的三个活性亚基被LMP取代，变成具有全新底物特异性和产物特异性的免疫蛋白酶体，其产物为常为6~30个氨基酸残基且C端多为疏水性或碱性氨基酸的多肽，更适合被内质网膜上的TAP（抗原相关转运物）转运至内质网腔</p>
</li>
<li><p>第二步 抗原肽转运至内质网</p>
</li>
<li><p>抗原肽与TAP结合后，TAP发生ATP依赖性蛋白构象改变，开放孔道。胞质内的抗原肽进入内质网腔，在与MHCI类分子结合过程中会被ERAP（抗原肽氨基肽酶）进一步修剪为8~12个氨基酸残基</p>
</li>
<li><p>TAP是一种异二聚体（TAP1&#x2F;2），TAP1、2各跨越内质网膜6次，形成孔样结构，依赖ATP对肽段进行主动转运</p>
</li>
<li><p>第三步 抗原肽-MHCI复合物组装</p>
</li>
<li><p>MHCI类分子的α链和β链在内质网中合成，Tapasin（TAP相关蛋白）介导新合成的MHCI类分子和TAP结合，在伴侣蛋白的作用下，MHCI类分子组装为二聚体，其α1&#x2F;2功能区构成抗原肽结合槽，与修剪后的抗原肽结合形成复合物，复合物与Tapasin解离</p>
</li>
<li><p>伴侣蛋白参与α链的折叠，α&#x2F;β链的组装；Erp57（羟基氧化还原酶）催化MHCI类分子功能区二硫键的断裂和重建，使抗原肽更加稳定</p>
</li>
<li><p>第四步 抗原肽-MHCI类分子复合物在细胞表面表达</p>
</li>
<li><p>复合物经高尔基体转运至细胞表面，等待CD8+T细胞识别</p>
</li>
</ul>
<h2 id="CTL细胞的产生"><a href="#CTL细胞的产生" class="headerlink" title="CTL细胞的产生"></a>CTL细胞的产生</h2><p>CD8+T细胞在CD8的协助下，通过αβTCR识别抗原肽-MHCI分子复合物，通过与<a target="_blank" rel="noopener" href="https://limour.top/1722.html">Th细胞活化</a>类似的方式，激活成CTL细胞</p>
<ul>
<li>CD8分子：αβ链通过二硫键形成的异源二聚体，</li>
<li>2条肽链均为跨膜蛋白，胞膜外区各含1个Ig样结构，能与MHCI类分子重链的α3结构域结合</li>
</ul>
<p>如果病原体直接感染专职APC细胞，则直接刺激CD8＋T细胞产生IL-2，诱导CTL细胞自身增殖分化为CTL</p>
<p>否则，在病灶区被第一信号激活的CD8＋T细胞需要Th1细胞分泌的IL2和被Th1的CD40L上调的APC的B7的第二信号活化。</p>
<h2 id="CTL细胞的效应"><a href="#CTL细胞的效应" class="headerlink" title="CTL细胞的效应"></a>CTL细胞的效应</h2><p>与Th1类似，CTL的归巢表型为选择素配体、CXCR3、CCR5</p>
<p>迁移至感染部位的CTL主要通过特异性识别表达抗原肽-MHCI类分子复合物的靶细胞（主要为病毒感染的细胞或肿瘤细胞），分泌和上调表达多种细胞杀伤分子，导致靶细胞凋亡，且杀伤过程中CTL自身不产生伤害，可连续对多个靶细胞进行杀伤。</p>
<ul>
<li>效-靶细胞结合：CTL高表达黏附分子并特异性识别抗原肽-MHCI类分子复合物，同时形成免疫突触浓集效应分子（为不影响邻近的正常细胞）</li>
<li>CTL极化：免疫突触形成后，CTL细胞膜分子&#x2F;包内成分聚集于细胞一端，对靶细胞发动致死性攻击</li>
<li>致死性攻击：<br>穿孔素&#x2F;颗粒酶：穿孔素使细胞坏死，颗粒酶促进细胞凋亡<br><a target="_blank" rel="noopener" href="https://zhuanlan.zhihu.com/p/42772317">死亡受体</a>：FasL和靶细胞表面的Fas结合启动FasL介导的外部死亡受体途径的细胞凋亡；TNF-α与TNFR1结合后诱导TNFR1的DD聚集募集衔接蛋白TRADD，TRADD可募集TRAF2、RIP和FADD等信号分子。TRAF2和RIP可激活NF-κB和JNK&#x2F;AP信号通路，而FADD可以激活Caspase级联反应，衔接蛋白TRADD募集信号分子的差异，决定了细胞的生存或死亡。</li>
<li>之后脱离靶细胞，寻找下一个目标</li>
<li>CTL也分泌细胞因子参与免疫防御</li>
</ul>

                
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