外泌体是由多数细胞分泌的细胞外微囊泡（EMV），参与细胞间通讯。它们存在于体液中，所含蛋白质种类繁多，取决于宿主细胞类型。其成分还会随细胞状态（如应激、激活或特定信号通路抑制）而变化，这使得外泌体成为癌症诊断等领域液体活检的理想生物标志物。
外泌体生物发生始于早期内体的内陷。这些内体成熟为晚期内体，进一步形成多泡体（MVBs）。MVB形成涉及内体膜向内出芽，导致细胞质成分被隔离至腔内囊泡（ILVs）中。ILVs通过胞吐作用释放到细胞外空间，外泌体因此得名。外泌体所含蛋白质标记物种类广泛，取决于宿主细胞及其状态（如应激或激活，或特定信号通路抑制）。四跨膜蛋白（如CD9、CD63和CD81）是最常见的规范性外泌体标记蛋白。四跨膜蛋白的表面定位使其特别适用于生物样本中外泌体的免疫标记和纯化。运输所需内体分选复合物（ESCRT）组分（如TSG101和Alix）、细胞骨架蛋白、整合素和膜联蛋白在外泌体上也高度富集。这些分子在外泌体靶向和细胞黏附中发挥关键作用。
什么是外泌体？
外泌体是小（50-120nm）的内体来源细胞外微囊泡（EMV），通过在细胞间运输蛋白质、脂质和核酸等生物分子，在细胞间通讯中起关键作用。它们最早于1980年代初在网织红细胞培养基中被观察到。"外泌体"一词源于其通过胞吐作用释放至细胞外介质的现象。外泌体具有与质膜相似的拓扑结构，几乎由所有细胞类型分泌，并已在所有体液中得到证实。
外泌体如何形成？
外泌体生物发生通常始于通过内吞作用对细胞外物质的内化，形成早期内体（一种包含内化货物的膜结合细胞器）。随后，早期内体成熟为晚期内体，此过程伴随膜成分变化和货物分选。晚期内体可发育为多泡体（MVBs），其特征是存在通过内体膜内陷形成的腔内囊泡（ILVs）。在MVB中，蛋白质、脂质、核酸和其他分子被分选至ILVs内。这是外泌体生物发生的关键步骤，由运输所需内体分选复合物（ESCRT）实现。ESCRT包含多个蛋白复合物（ESCRT-0、ESCRT-I、ESCRT-II、ESCRT-III），它们协同工作以识别并将货物隔离至出芽的ILVs中。MVBs随后可能与溶酶体融合降解，或与质膜融合，将ILVs作为外泌体释放至细胞外空间。
如何识别外泌体？
外泌体所含蛋白质种类繁多，取决于其宿主细胞，且其成分受细胞状态（如应激、激活或特定信号通路抑制）调控。四跨膜蛋白（如CD9、CD63和CD81）是最常见的规范性外泌体标记蛋白，位于囊泡表面。四跨膜蛋白抗原的表面定位使其成为生物样本中外泌体免疫标记和纯化的理想靶标。运输所需内体分选复合物（ESCRT）组分（如TSG101和Alix）、细胞骨架蛋白、整合素和膜联蛋白在外泌体上也高度富集；这些分子在外泌体靶向和细胞黏附中起关键作用。
外泌体为何重要？
外泌体分泌是持续进行的，尽管其分泌速率和组成可能受多种内在或外在因素（如细胞应激、信号级联）影响。尽管普遍存在，外泌体被视为非经典分泌途径组分。
由于外泌体几乎由所有细胞类型分泌，其组成反映了宿主的多样性，并高度依赖于起源细胞类型。其分子组成也反映起源细胞或组织的生理或病理生理变化。这使得外泌体成为液体活检的优异生物标志物，用于诊断和追踪疾病进展。在外泌体诊断中，相较于循环肿瘤细胞（CTCs）或循环肿瘤DNA（ctDNA），外泌体具有显著优势：丰度高、稳定性强、所含标志物分子种类广泛。外泌体还参与细胞间通讯。外泌体组分可能直接转移至邻近细胞，或通过称为转胞吞作用的方法在不同细胞间穿梭后到达最终目的地。通过这种方式，外泌体可在简单扩散不足的远距离传递信号。其在细胞间通讯中的作用表明，外泌体可能在许多生理过程中具有更深层次角色；这一假设得到以下观察支持：外泌体信号在发育与模式形成、免疫反应、神经元通讯和组织修复中发挥直接作用。在某些病理中，外泌体也充当载体；肿瘤细胞衍生的外泌体在肿瘤血管生成和转移中发挥主动作用。受刺激的血细胞和血管内皮细胞脱落的外泌体参与多发性硬化症、短暂性脑缺血发作和抗磷脂综合征等神经系统疾病。外泌体还可能携带靶向破坏的受损细胞物质，并促进α-突触核蛋白和β-淀粉样蛋白等聚集蛋白的毒性形式扩散，从而推动神经退行性疾病进展。一些研究还表明，外泌体运输已被SARS-CoV-2等病毒病原体利用，在宿主细胞间传播并逃避免疫检测。由于其体积小、结构简单，外泌体有时可穿过血脑屏障。有观点认为，外泌体可作为靶向中枢神经系统的递送系统，在无需侵入性手术的情况下治疗神经病理性疾病。利用外泌体转移遗传信息或递送治疗剂，是一个目前尚未充分探索但具有巨大医学潜力的领域。
如何研究外泌体？
外泌体分泌组庞大且多样，包含多种不同标记物（参见exocarta.org）。上述规范性表面标记物的存在，使得从不同类型样本中纯化和深入研究外泌体分泌及内容成为可能。
外泌体标记蛋白抗体
根据近期文献，最相关的蛋白质外泌体标记物包括：
| 蛋白质 | 基因 | 基因ID | UniProt | 参考 | exocarta前100蛋白 | TS | EF | LP | TA | CS | AG | MT | AP | HS | EN | RG | CA | II | VI | ND |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| 14-3-3蛋白ε | YWHAE | 7531 | P62258 | - | 22 | X | ||||||||||||||
| 14-3-3蛋白β/α | YWHAB | 7529 | P31946 | 50 | x | |||||||||||||||
| 14-3-3蛋白η | YWHAH | 7533 | Q04917 | 94 | x | |||||||||||||||
| 14-3-3蛋白γ | YWHAG | 7532 | P61981 | 54 | x | |||||||||||||||
| 14-3-3蛋白θ | YWHAQ | 10971 | P27348 | 56 | x | |||||||||||||||
| 14-3-3蛋白ζ/δ | YWHAZ | 7534 | P63104 | - | 15 | X | ||||||||||||||
| 78kDa葡萄糖调节蛋白 | HSPA5 | 3309 | P11021 | (1) | 35 | X | ||||||||||||||
| 肌动蛋白，胞质1型 | ACTB | 60 | P60709 | - | 5 | X | ||||||||||||||
| ADAM10 | ADAM10 | 102 | O14672 | (2) | - | X | X | X | ||||||||||||
| Alix | PDCD6IP | 10015 | Q8WUM4 | (3) | 2 | X | ||||||||||||||
| α-烯醇化酶 | ENO1 | 2023 | P06733 | (4) | 9 | X | ||||||||||||||
| α-突触核蛋白 | SNCA | 6622 | P37840 | (32) | - | X | X | |||||||||||||
| 氨肽酶N | ANPEP | 290 | P15144 | - | - | X | X | |||||||||||||
| β-淀粉样蛋白 | APP | 351 | P05067 | (5) | - | X | ||||||||||||||
| 膜联蛋白A1 | ANXA1 | 301 | P04083 | 53 | x | |||||||||||||||
| 膜联蛋白A11 | ANXA11 | 311 | P50995 | 68 | x | |||||||||||||||
| 膜联蛋白A4 | ANXA4 | 307 | P09525 | 72 | x | |||||||||||||||
| 膜联蛋白A6 | ANXA6 | 309 | P08133 | 67 | x | |||||||||||||||
| 膜联蛋白A5 | ANXA5 | 308 | P08758 | (6) | 20 | X | X | |||||||||||||
| 膜联蛋白A2 | ANXA2 | 302 | P07355 | (7) | 6 | X | ||||||||||||||
| AP-1 | JUN | 3725 | P05412 | - | - | X | X | |||||||||||||
| 三磷酸柠檬酸裂解酶 | ACLY | 47 | P53396 | - | 72 | X | ||||||||||||||
| ATP酶 | ATP1A1 | 476 | P05023 | - | 39 | X | ||||||||||||||
| 基质金属蛋白酶诱导蛋白 | BSG | 682 | P35613 | (8) | - | X | X | |||||||||||||
| 小窝蛋白-1 | CAV1 | 857 | Q03135 | (9), (10) | - | X | X | |||||||||||||
| CD9 | CD9 | 928 | P21926 | (11) | 1 | X | ||||||||||||||
| CD11a | ITGAL | 3683 | P20701 | (12) | - | X | X | |||||||||||||
| CD11b | ITGAX | 3687 | P11215 | (12) | - | X | X | |||||||||||||
| CD11c | ITGAM | 3684 | P20702 | (12) | - | X | X | |||||||||||||
| CD29 | ITGB1 | 3688 | P05556 | (12) | 34 | X | X | |||||||||||||
| CD37 | CD37 | 951 | P11049 | (11) | - | X | ||||||||||||||
| CD44 | CD44 | 960 | P16070 | (13) | - | X | X | |||||||||||||
| CD49f | ITGA6 | 3655 | P23229 | (12) | 89 | X | X | |||||||||||||
| CD53 | CD53 | 963 | P19397 | - | x | |||||||||||||||
| CD63 | CD63 | 967 | P08962 | (10), (11) | 7 | X | X | |||||||||||||
| CD81 | CD81 | 975 | P60033 | (14) | 24 | X | X | |||||||||||||
| CD82 | CD82 | 3732 | P27701 | (11) | - | X | ||||||||||||||
| CD142 | TF | 2152 | P13726 | (15) | - | X | X | |||||||||||||
| CD146 | MCAM | 4162 | P43121 | (15) | - | X | X | |||||||||||||
| CD163 | CD163 | 9332 | Q86VB7 | (15) | - | X | X | X | ||||||||||||
| 网格蛋白重链1 | CLTC | 1213 | Q00610 | - | 23 | X | ||||||||||||||
| 闭合蛋白-1 | CLDN1 | 9076 | O95832 | (8) | - | X | ||||||||||||||
| 丝切蛋白-1 | CFL1 | 1072 | P23528 | - | 25 | X | ||||||||||||||
| EBV LMP1 | - | - | - | (16) | - | X | ||||||||||||||
| EBV LMP2A | - | - | - | (16) | - | X | ||||||||||||||
| EF-1-α-1 | EEF1A1 | 1915 | P68104 | (4) | 14 | X | ||||||||||||||
| EF2 | EEF2 | 1938 | P13639 | - | 17 | X | ||||||||||||||
| EGFR | EGFR | 1956 | P00533 | (15) | - | X | ||||||||||||||
| Ep-CAM | EPCAM | 4072 | P16422 | (17), (18) | - | X | X | |||||||||||||
| 脂肪酸合酶 | FASN | 2194 | P49327 | (3) | 21 | X | X | |||||||||||||
| 纤连蛋白 | FN1 | 2335 | P02751 | 93 | ||||||||||||||||
| 上浮蛋白-1 | FLOT1 | 10211 | O75955 | (18), (19) | 41 | X | X | |||||||||||||
| 上浮蛋白-2 | FLOT2 | 2319 | Q14254 | (19) | - | X | ||||||||||||||
| 果糖-1,6-二磷酸醛缩酶A | ALDOA | 226 | P04075 | - | 18 | X | ||||||||||||||
| 凝溶胶蛋白 | GSN | 2934 | P06396 | 92 | x | x | ||||||||||||||
| 甘油醛-3-磷酸脱氢酶 | GAPDH | 2597 | P04406 | - | 4 | X | ||||||||||||||
| HCV核心蛋白 | - | - | - | (20) | - | X | ||||||||||||||
| 热休克70kDa蛋白1A | HSPA1A | 3303 | P0DMV8 | 51 | x | |||||||||||||||
| 热休克蛋白HSP 90-α | HSP90AA1 | 3320 | P07900 | (1) | 10 | X | X | |||||||||||||
| 热休克蛋白HSP 90-β | HSP90AB1 | 3326 | P08238 | (1) | 19 | X | ||||||||||||||
| 肝素酶 | HPSE | 10855 | Q9Y251 | (21) | - | X | X | |||||||||||||
| HIV Gag | - | - | - | (20) | - | X | ||||||||||||||
| HIV Nef | - | - | - | (20) | - | X | ||||||||||||||
| HLA-DRA | HLA-DRA | 3122 | P01903 | (22) | - | X | X | X | X | |||||||||||
| HLA-G | HLA-G | 3135 | P17693 | (23) | - | X | X | X | X | |||||||||||
| Hsc70 | HSPA8 | 3312 | P11142 | (1) | 3 | X | ||||||||||||||
| HSV gB | - | - | - | (16) | - | X | ||||||||||||||
| HTLV-1 Tax | Tax | - | - | (20) | - | X | ||||||||||||||
| 亨廷顿蛋白 | HTT | 3064 | P42858 | (5) | - | X | X | |||||||||||||
| ICAM-1 | ICAM1 | 3383 | P05362 | (24) | - | X | ||||||||||||||
| 富亮氨酸受体激酶2 | LRRK2 | 120892 | Q5S007 | (5) | - | X | ||||||||||||||
| L-乳酸脱氢酶A链 | LDHA | 3939 | P00338 | - | 13 | X | ||||||||||||||
| 溶酶体相关膜糖蛋白1 | LAMP1 | 3916 | P11279 | (25) | - | X | X | |||||||||||||
| 溶酶体相关膜糖蛋白2 | LAMP2 | 3920 | P13473 | (23) | 88 | X | X | |||||||||||||
| MHCI | - | - | - | (26) | - | X | X | |||||||||||||
| MHCII | - | - | - | (26) | - | X | X | |||||||||||||
| MUC1 | MUC1 | 4582 | P15941 | (15) | - | X | X | |||||||||||||
| N-钙黏蛋白 | CDH2 | 1000 | P19022 | (15) | - | X | X | |||||||||||||
| 磷酸甘油酸激酶1 | PGK1 | 5230 | P00558 | (4) | 16 | X | ||||||||||||||
| 胎盘碱性磷酸酶 | ALPP | 250 | P05187 | (15) | - | X | X | |||||||||||||
| 朊病毒蛋白 | - | - | - | (5) | - | X | ||||||||||||||
| 前列腺特异性抗原 | KLK3 | 354 | P07288 | (27) | - | X | X | |||||||||||||
| 丙酮酸激酶PKM | PKM | 5315 | P14618 | (28) | 12 | X | X | |||||||||||||
| Rab-14 | RAB14 | 51552 | P61106 | - | 75 | X | ||||||||||||||
| Rab-5a | RAB5A | 5868 | P20339 | - | 80 | X | ||||||||||||||
| Rab-5b | RAB5B | 5869 | P61020 | - | 86 | X | ||||||||||||||
| Rab-5c | RAB5C | 5878 | P51148 | - | 64 | X | ||||||||||||||
| Rab-7 | RAB7A | 7879 | P51149 | - | 61 | X | ||||||||||||||
| Rap 1B | RAP1B | 5908 | P61224 | - | 33 | X | ||||||||||||||
| 超氧化物歧化酶 | SOD1 | 6647 | P00441 | - | x | x | ||||||||||||||
| 乙酰肝素硫酸蛋白聚糖-1 | SDC1 | 6382 | P18827 | (29) | - | X | ||||||||||||||
| 乙酰肝素硫酸蛋白聚糖-4 | SDC4 | 6385 | P31431 | (29) | - | X | ||||||||||||||
| Syntenin-1 | SDCBP | 6386 | O00560 | (30) | 8 | X | ||||||||||||||
| TARDBP | TDP-43 | 23435 | Q13148 | - | x | |||||||||||||||
| 内质网ATP酶 | VCP | 7415 | P55072 | 26 | ||||||||||||||||
| 三磷酸甘油醛异构酶 | TPI1 | 7167 | P60174 | 27 | x | |||||||||||||||
| 肿瘤相关糖蛋白 | TAG-72 | - | - | (15) | - | X | ||||||||||||||
| 四跨膜蛋白-8 | Tspan8 | 7103 | P19075 | (15) | - | X | X | |||||||||||||
| Tsg101 | TSG101 | 7251 | Q99816 | (31) | 11 | X | ||||||||||||||
| 微管蛋白α-1C链 | TUBA1C | 84790 | Q9BQE3 | - | x | |||||||||||||||
| 微管蛋白α-4A链 | TUBA4A | 7277 | P68366 | - | x | |||||||||||||||
| 微管蛋白β-2B链 | TUBB2B | 347733 | Q9BVA1 | - | x | |||||||||||||||
| 微管蛋白β-4B链 | TUBB4B | 10383 | P68371 | - | x | |||||||||||||||
| 液泡分选蛋白35 | VPS35 | 55737 | Q96QK1 | (5) | - | X | X |
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