吞噬細胞

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長桿形的細菌,其中一個已經被一個稍大的球狀白血球部分吞噬。白血球因爲其體內尚未消化的細菌體而變形
一張中型粒細胞吞噬炭疽桿菌(橘黃色)的掃描電鏡照片

吞噬細胞(Phagocytes)为一类通过吞噬诸如细菌、坏死凋亡的细胞这样的有害外来粒子来保护机体的细胞。其英文名「Phagocytes」的前一部分来自希腊语「phagein」(意为食用、吞食),后一部分「-cyte」为生物学中表示细胞(cell)的词缀,来自于希腊语「kutos」(意为「中空容器」)[1]。吞噬细胞在对抗感染以及後續的免疫过程中不可或缺[2]吞噬细胞在整个动物界中都相当重要[3],且在脊椎动物体内高度发达[4]。一公升的人类血液约含六十亿个吞噬细胞[5]。1882年,埃黎耶·埃黎赫·梅契尼可夫在研究海星幼虫时发现了吞噬细胞[6]。他因这一成就获得了1908年的诺贝尔生理医学奖[7]许多物种体内都有吞噬细胞。一些阿米巴的行为亦与巨噬细胞(吞噬细胞的一种)相似,说明吞噬细胞在生命进化的早期阶段就出现了[8]

人和其他动物的吞噬细胞可由他们在吞噬过程中的效果分为专职(professional)吞噬细胞和非专职(non-professional)吞噬细胞[9]。许多类型的白血球都是专职吞噬细胞(如嗜中性粒細胞單核細胞巨噬細胞肥大細胞樹突狀細胞等)[10]。专职吞噬细胞和非专职吞噬细胞的主要区别在于专职吞噬细胞表面有一种叫受体的分子。受体可以探测细菌这样的在一般情况下不会在体内出现的有害物质[11]。吞噬细胞在对抗感染的过程中发挥很大的作用。吞噬细胞亦能够清除坏死或凋亡的细胞,使生物的各组织保持健康状态[12]

在感染过程中,化学信号使吞噬细胞前往病原体侵入机体处。这些化学物质可能来自于细菌,也可能来自其它已经出现的吞噬细胞。吞噬细胞通过趋化作用进行移动。当吞噬细胞与细菌接触后,吞噬细胞表面的受体将会与细菌结合。这样的结合将介导吞噬细胞将细菌吞噬[13]。一些吞噬细胞隨後通过活性氧類(ROS)以及一氧化氮杀死被它吞噬的细菌[14]。在吞噬完成后,巨噬细胞以及树突状细胞亦可参与抗原呈递过程。在该过程中,吞噬细胞将被它吞噬的物质的一部分转运到它的表面。随后,这些物质会被展示给其它的免疫细胞。一些吞噬细胞会移动到生物体的淋巴结处,并将上述物质呈递给一种白血球,淋巴细胞。该过程对建立免疫相当重要[15]。不过,许多病原体都进化出了逃避吞噬细胞攻击的机制[2]

歷史[编辑]

一個大鬍子老人拿着一根試管。他坐在一張窗邊的椅子上。桌上堆滿了小瓶子和試管
埃黎耶·埃黎赫·梅契尼可夫在他的實驗室裏

俄羅斯動物學家埃黎耶·埃黎赫·梅契尼可夫(1845–1916)爲第一個發現了吞噬細胞這種參與肌體對抗微生物感染的特化細胞的人。1882年,他在研究海星幼虫体内的能动(自由移动)细胞时,相信这类细胞对动物的免疫防御相当重要。为了验证他的猜想,他将一段取自柑橘树的小木刺插入了海星体内。几个小时后,他注意到,能动细胞聚集在了木刺周围[16]。梅契尼可夫之后去了维也纳,和卡爾·弗里德里希·威廉·克勞斯英语Carl Friedrich Wilhelm Claus(Carl Friedrich Wilhelm Claus)分享他的想法。卡尔提议,将梅契尼可夫发现的细胞命名为吞噬细胞(「phagocyte」)[17]。吞噬细胞(phagocyte)这个名字来源于希腊语单词「phagein」(吃,吞食)和「kutos」(中空容器)[1]

一年之后,梅契尼可夫对一种淡水甲壳动物水蚤进行了研究。水蚤是一种微小的、通体透明的动物,可在显微镜下对其进行观察。梅契尼可夫发现攻击水蚤的真菌都被水蚤的吞噬细胞消灭。随后,他又对哺乳动物的白血球进行观察,发现吞噬细胞通过一种被他称为吞噬(phagocytosis)的过程吞食消灭掉炭疽桿菌Bacillus anthracis,一种细菌)[18]。梅契尼可夫随后提出,吞噬细胞为机体针对入侵生物的防御体系的重要组成部分。

1903年,阿尔姆罗思·赖特英语Almroth Wright发现,特殊的抗体(他称之为调理素英语Opsonin(opsonin)。调理素(Opsonin)这个单词来自希腊语单词「opson」(意为调味品))可增强吞噬过程[19]。梅契尼可夫因为他对吞噬细胞和吞噬的研究,在1908年和保罗·埃尔利希共享了诺贝尔生理医学奖[7]

尽管人们在20世纪早些时候逐渐意识到这些发现的重要性,但吞噬细胞与免疫系统其他元件之间错综复杂直到1980年代才被阐明[20]

吞噬作用[编辑]

吞噬过程分三步:第一步,未与相关信号分子结合的吞噬细胞表面受体不会引发吞噬作用;第二步,受体与相关信号分子结合后,聚成簇状;第三步,吞噬作用引发,微粒被吞噬细胞吞入胞内

吞噬(Phagocytosis)指细胞吞食细菌、寄生虫死亡细胞等颗粒,以及细胞残骸、外来的碎屑的过程[21]。该过程涉及一系列的连续的分子过程[22]。吞噬在异物(这里以细菌为例)与吞噬细胞表面一种名为「受体」(receptors)的分子结合后发生。吞噬细胞随后运动到细菌周围,并将之吞食。人中性粒細胞对细菌进行吞噬平均要花9分钟[23]。被吞噬细胞吞入胞内的细菌会被裹在吞噬小体内。在一分钟内,吞噬小体即会与一个溶酶体或一个顆粒英语Granule (cell biology)融合,形成一个吞噬溶酶體英语phagolysosome。细菌随后会暴露在超量的杀灭性物质之中[24],並於幾分鐘後死亡[23]。树突状细胞以及巨噬细胞的吞噬速度相对较慢,吞噬在这些细胞中可能会花掉数个小时。巨噬细胞是巨大而且有些「乱糟糟」的吞噬者。它们会吞噬许多的物质,偶尔还会将一些未消化的物质释放回组织之中。未消化的碎片起到一个信号的作用,它们能从血液中征募更多的吞噬细胞[25]。吞噬细胞相当「贪婪」。科学家曾经用铁屑来饲喂巨噬细胞,之后,他们用小磁铁从其他细胞中吸出了这些铁屑[26]

A cartoon: The macrophage is depicted as a distorted solid circle. On the surface of the circle is a small y-shaped figure that is connected to a solid rectangle that depicts a bacterium.
巨噬细胞有特殊的受体,参与能增强吞噬作用的过程(图中各物件未按照实际比例绘制)

进化起源[编辑]

吞噬作用在生物界中十分普遍,而且可能在进化的早期[123],即单细胞真核生物阶段就出现了[124]阿米巴是一类单细胞原生动物。植物从进化树上分出後不久,出现了阿米巴。阿米巴是後生動物的祖先。阿米巴与吞噬细胞共有许多特殊功能[124]。比如,盘基网柄菌Dictyostelium discoideum)是一种生活在土壤和草料中的阿米巴,它们以细菌为食。和动物的吞噬细胞一样,它通过柱形受体,经由吞噬过程吞食细菌。除此以外,它与吞噬细胞还有很多共同的生物学功能[125]。盘基网柄菌具有社会性。在饥饿时,它们会聚集在一起,生成一团假合胞(pseudoplasmodium)或蛞蝓形的物质(slug)英语Dictyostelid。这个多细胞集群能够形成一团有孢子子實體,来对抗环境危险。在子实体生成前,盘基网柄菌细胞会在几天里以一团蛞蝓形的物质迁移。与致病细菌或毒素接触可能会造成产孢受阻,使物种生存受到威胁。不过,一些在细胞集群中巡弋的阿米巴细胞会吞噬相关的细菌或吸收毒素,并在完成上述过程中死亡。它们与细胞集群中的其它阿米巴细胞在遗传学上完全相同。它们的自我牺牲保护了其它的阿米巴细菌不受细菌的侵害,这和高等脊椎动物的免疫系统中的吞噬细胞的自我牺牲行为相似。这一社会性阿米巴的原始免疫现象说明,一种进化上保守的细胞吞噬机制可能在阿米巴进化成更高等的生物前就被广泛用于防御[126]。然而,一个哺乳动物吞噬细胞的共同祖先还未被发现。吞噬细胞在整个动物界中都有分布[3],从海绵动物到昆虫,再到低等和高等脊椎动物体内,都有吞噬细胞[127][128]。阿米巴区别自我和非我的能力非常重要,是许多阿米巴的免疫系统的根源[8]

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