干细胞来源的胞外囊泡对T细胞的免疫调控作用

doi:10.3877/cma.j.issn.2095-1221.2019.04.011

胞外囊泡（EVs）是细胞旁分泌产生的一种亚细胞成分，实质上是一组纳米级颗粒。它是双层膜结合型囊泡，内含蛋白质、核酸等活性成分。EVs在细胞间通过转移携带的信号分子而获得重要的地位。目前关于EVs在体外和体内的研究中对T细胞的调控能力引起了人们广泛的兴趣。在大多数研究中干细胞被报道能够抑制T细胞的增殖、活化和分化，在极少数研究中也发现干细胞具有增强T细胞免疫反应的作用。事实上所有的细胞类型均能释放EVs，包括干/祖/前体细胞。EVs被认为是细胞间交流的一种新机制，具有与干/祖细胞等亲代细胞相似的免疫调控作用。本综述是概述干/祖细胞来源的EVs对T细胞调控作用及可能的机制。

关键词: 干细胞, 胞外囊泡, T细胞, 免疫调节

Extracellular vesicles (EVs) are subcellular component secreted by paracellular mechanism, which are essentially a group of nanoparticles. They are bilayer membrane bound vesicles containing proteins, nucleic acids and other active components. EVs gain significant status by the transfer of signal molecules among cells. Recent studies on the regulatory effects of EVs on T cells both in vitro and in vivo have aroused extensive interest. In most investigations, stem cells have been reported to suppress the proliferation, activation and differentiation of T cells, meanwhile, stem cells also have been found to enhance the immune response of T cells in few reports. Virtually all cell types have the ability to release EVs including stem/progenitor cells. EVs are recognized as a novel mechanism of intercellular communication, with immunoregulatory effects similar to those of parental cells such as stem/progenitor cells. The aim of this review is to summarize the possible mechanisms underlying the regulatory effects of stem/progenitor cell-derived EVs on T cells.

Key words: Stem cell, Extracellular vesicle, T cell, Immune regulation

表1 干细胞来源的胞外囊泡对T细胞的免疫调控作用

释放EVs的干细胞类型	模型	转移物质	靶细胞
hBM-MSCs	T1D患者	PGE2和TGF-β	T细胞
hBM-MSCs	体外	—	T细胞
mAD-MSCs	T1D小鼠模型	—	T细胞
hPDL-MSCs（LPS pretreated）	慢性牙周炎	microRNA-155-5p	T细胞
hNSCs	血栓性卒中小鼠模型	—	巨噬细胞，T细胞
hAD-MSCs	实验性过敏性哮喘小鼠模型	—	T细胞
hBM-MSCs	严重的顽固性哮喘小鼠模型	—	T细胞
hBM-MSCs	体外	—	T细胞
mBM-MSCs	炎性关节炎小鼠模型	—	T细胞
hUC-MSCs	GVHD小鼠模型	—	T细胞
iPSC-MSCs（Protein-free medium activated）	干燥综合征小鼠模型	—	APCs，T细胞
hBM-MSCs	T1D患者	—	DCs，T细胞
hMSCs（Protein-free medium activated）	T1D以及葡萄膜视网膜炎小鼠模型	—	DCs，T细胞
hGSCs	体外	—	CD14⁺单核细胞，T细胞
hESC-MSCs	同种异体皮肤移植物小鼠模型	TLL4	单核细胞，T细胞
mBM-MSCs	系统性硬化小鼠模型	miR-151-5p	受者BM-MSCs，Th2细胞
hBM-MSCs	aGVHD小鼠模型	miR-125a-3p	T细胞
hUC-MSCs	体外	CD73蛋白	T细胞
canine WJ-MSCs	体外	TGF-β和腺苷信号	T细胞
h-endMSCs	体外	TGF-β	T细胞
mESC	移植的肺腺癌小鼠模型	表达GM-CSF	CD8⁺ T细胞，Tregs细胞
rMSC	肝细胞癌大鼠模型	β-catenin	NK-T细胞
hBM-MSCs	卒中大鼠、小鼠模型	—	T细胞，B细胞，NK细胞
mAD-SC	实验性自身免疫性脑脊髓膜炎小鼠模型	—	T细胞
hUC-MSC	实验性自身免疫性葡萄膜炎大鼠模型	—	视网膜细胞
hBM-MSC	人-鼠异种GVHD模型	腺苷信号	Th1细胞
释放EVs的干细胞类型	生物机制	生物作用	参考文献
hBM-MSCs	调控Th17/Tregs免疫反应轴	阻止疾病进展	[7]
hBM-MSCs	同[7]	诱导免疫耐受	[17]
mAD-MSCs	同[7]	阻止疾病进展	[18]
hPDL-MSCs （LPS pretreated）	靶向sirtuin-1以及调控Th17/Tregs免疫反应轴	阻止牙周炎的进一步恶化	[19]
hNSCs	同[7]	治疗作用以及改善预后	[20]
hAD-MSCs	将Th2细胞的免疫反应向Th1转变	抑制炎症和组织重塑	[21]
hBM-MSCs	将Th2/Th17细胞的免疫反应向Th1转变	减弱气道炎症	[22]
hBM-MSCs	诱导Th1的免疫反应向Th2细胞转换	调控T细胞的免疫反应	[17]
mBM-MSCs	Th1细胞的免疫反应向Th2变换	治疗关节炎	[23]
hUC-MSCs	抑制细胞毒性T细胞，将Th1细胞的免疫反应向Th2转变	阻止致命的GVHD	[24]
iPSC-MSCs（Protein-free medium activated）	抑制Tfh/Th17细胞分化	阻止疾病进展	[25]
hBM-MSCs	诱导DCs未成熟表型，抑制Th1/Th17细胞分化，增加Tregs细胞	诱导免疫耐受	[26]
hMSCs（Protein-free medium activated）	诱导DCs未成熟表型以及抑制Th1/Th17细胞发育	减弱炎性反应	[27]
hGSCs	转变单核细胞表型以及抑制T细胞活化、分化和增殖	诱导免疫耐受	[29]
hESC-MSCs	调控单核细胞朝向M2表型转变以及调控CD4⁺ T细胞向Tregs分化	促进皮肤存活	[30]
mBM-MSCs	抑制IL4Rα/ mTOR路径以及抑制Th2细胞活化和浸润	系统性硬化的治疗靶点	[31]
hBM-MSCs	保存循环初始T细胞	延长aGVHD小鼠的生存期	[35]
hUC-MSCs	抑制T细胞增殖	免疫抑制反应	[39]
canine WJ-MSCs	抑制CD4⁺ T细胞增殖	免疫调控作用	[42]
h-endMSCs	抑制CD4⁺ T细胞活化	免疫调控作用	[44]
mESC	降低瘤内Tregs，增加瘤内CD8⁺ T/Tregs比率，激活瘤内CD8⁺ CD25⁺、CD8⁺ IFN-γ⁺效应细胞	用于阻止肺癌的预防性疫苗	[45]
rMSC	促进NK-T细胞存活以及促进NK-T细胞向瘤内迁移	促进NK-T细胞的抗肿瘤反应以及导致低级别肿瘤分化	[46]
hBM-MSCs	削弱T细胞、B细胞、NK细胞淋巴细胞减少症	阻止缺血后的免疫抑制	[48,49]
mAD-SC	抑制整合素依赖趋化因子诱导的路径以及抑制活化的T细胞黏附	减弱慢性炎症	[50]
hUC-MSC	下调CCL21表达以及减少T细胞浸润	保护视网膜免于炎性损伤	[51]
hBM-MSC	诱导Th1细胞凋亡	促进免疫抑制	[43]

表1 干细胞来源的胞外囊泡对T细胞的免疫调控作用

释放EVs的干细胞类型	模型	转移物质	靶细胞
hBM-MSCs	T1D患者	PGE2和TGF-β	T细胞
hBM-MSCs	体外	—	T细胞
mAD-MSCs	T1D小鼠模型	—	T细胞
hPDL-MSCs（LPS pretreated）	慢性牙周炎	microRNA-155-5p	T细胞
hNSCs	血栓性卒中小鼠模型	—	巨噬细胞，T细胞
hAD-MSCs	实验性过敏性哮喘小鼠模型	—	T细胞
hBM-MSCs	严重的顽固性哮喘小鼠模型	—	T细胞
hBM-MSCs	体外	—	T细胞
mBM-MSCs	炎性关节炎小鼠模型	—	T细胞
hUC-MSCs	GVHD小鼠模型	—	T细胞
iPSC-MSCs（Protein-free medium activated）	干燥综合征小鼠模型	—	APCs，T细胞
hBM-MSCs	T1D患者	—	DCs，T细胞
hMSCs（Protein-free medium activated）	T1D以及葡萄膜视网膜炎小鼠模型	—	DCs，T细胞
hGSCs	体外	—	CD14⁺单核细胞，T细胞
hESC-MSCs	同种异体皮肤移植物小鼠模型	TLL4	单核细胞，T细胞
mBM-MSCs	系统性硬化小鼠模型	miR-151-5p	受者BM-MSCs，Th2细胞
hBM-MSCs	aGVHD小鼠模型	miR-125a-3p	T细胞
hUC-MSCs	体外	CD73蛋白	T细胞
canine WJ-MSCs	体外	TGF-β和腺苷信号	T细胞
h-endMSCs	体外	TGF-β	T细胞
mESC	移植的肺腺癌小鼠模型	表达GM-CSF	CD8⁺ T细胞，Tregs细胞
rMSC	肝细胞癌大鼠模型	β-catenin	NK-T细胞
hBM-MSCs	卒中大鼠、小鼠模型	—	T细胞，B细胞，NK细胞
mAD-SC	实验性自身免疫性脑脊髓膜炎小鼠模型	—	T细胞
hUC-MSC	实验性自身免疫性葡萄膜炎大鼠模型	—	视网膜细胞
hBM-MSC	人-鼠异种GVHD模型	腺苷信号	Th1细胞
释放EVs的干细胞类型	生物机制	生物作用	参考文献
hBM-MSCs	调控Th17/Tregs免疫反应轴	阻止疾病进展	[7]
hBM-MSCs	同[7]	诱导免疫耐受	[17]
mAD-MSCs	同[7]	阻止疾病进展	[18]
hPDL-MSCs （LPS pretreated）	靶向sirtuin-1以及调控Th17/Tregs免疫反应轴	阻止牙周炎的进一步恶化	[19]
hNSCs	同[7]	治疗作用以及改善预后	[20]
hAD-MSCs	将Th2细胞的免疫反应向Th1转变	抑制炎症和组织重塑	[21]
hBM-MSCs	将Th2/Th17细胞的免疫反应向Th1转变	减弱气道炎症	[22]
hBM-MSCs	诱导Th1的免疫反应向Th2细胞转换	调控T细胞的免疫反应	[17]
mBM-MSCs	Th1细胞的免疫反应向Th2变换	治疗关节炎	[23]
hUC-MSCs	抑制细胞毒性T细胞，将Th1细胞的免疫反应向Th2转变	阻止致命的GVHD	[24]
iPSC-MSCs（Protein-free medium activated）	抑制Tfh/Th17细胞分化	阻止疾病进展	[25]
hBM-MSCs	诱导DCs未成熟表型，抑制Th1/Th17细胞分化，增加Tregs细胞	诱导免疫耐受	[26]
hMSCs（Protein-free medium activated）	诱导DCs未成熟表型以及抑制Th1/Th17细胞发育	减弱炎性反应	[27]
hGSCs	转变单核细胞表型以及抑制T细胞活化、分化和增殖	诱导免疫耐受	[29]
hESC-MSCs	调控单核细胞朝向M2表型转变以及调控CD4⁺ T细胞向Tregs分化	促进皮肤存活	[30]
mBM-MSCs	抑制IL4Rα/ mTOR路径以及抑制Th2细胞活化和浸润	系统性硬化的治疗靶点	[31]
hBM-MSCs	保存循环初始T细胞	延长aGVHD小鼠的生存期	[35]
hUC-MSCs	抑制T细胞增殖	免疫抑制反应	[39]
canine WJ-MSCs	抑制CD4⁺ T细胞增殖	免疫调控作用	[42]
h-endMSCs	抑制CD4⁺ T细胞活化	免疫调控作用	[44]
mESC	降低瘤内Tregs，增加瘤内CD8⁺ T/Tregs比率，激活瘤内CD8⁺ CD25⁺、CD8⁺ IFN-γ⁺效应细胞	用于阻止肺癌的预防性疫苗	[45]
rMSC	促进NK-T细胞存活以及促进NK-T细胞向瘤内迁移	促进NK-T细胞的抗肿瘤反应以及导致低级别肿瘤分化	[46]
hBM-MSCs	削弱T细胞、B细胞、NK细胞淋巴细胞减少症	阻止缺血后的免疫抑制	[48,49]
mAD-SC	抑制整合素依赖趋化因子诱导的路径以及抑制活化的T细胞黏附	减弱慢性炎症	[50]
hUC-MSC	下调CCL21表达以及减少T细胞浸润	保护视网膜免于炎性损伤	[51]
hBM-MSC	诱导Th1细胞凋亡	促进免疫抑制	[43]

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Regulatory effects of stem cell-derived extracellular vesicles on T-lymphocytes

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Regulatory effects of stem cell-derived extracellular vesicles on T-lymphocytes