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程序性细胞死亡和凋亡研究历史：细胞死亡史上的重要事件
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吞噬细胞（Phagocytes）是Ilya (Elie) Metchnikoff（梅契尼科夫）首先描述和记录的，他意识到吞噬细胞吞噬了死亡的细胞。到20世纪中叶，研究人员开始探索细胞是如何死亡的，已经认识到细胞死亡是一个生理控制的过程，最常见的死亡模式（“萎缩坏死”，后来的凋亡）受到严格控制，在1990年之前，一些发现阐明了细胞凋亡的机制。21世纪初，认为自噬过程与细胞凋亡有关，可以限制或增加细胞死亡。之后，研究转向了什么代谢或其他机制间接控制细胞死亡正反馈回路的激活或抑制的问题。
推动这一领域发展的是三项科学进步：
技术上的突破：DNA电泳技术可以证明细胞凋亡，因为在坏死细胞中随机降解的DNA会产生涂片，而在细胞凋亡中，DNA在核小体之间以更有序的方式切割。
概念上的突破：认识到一些疾病，特别是癌症，与细胞死亡的异常模式有关。
理论上的突破：认识到细胞死亡的机制，甚至细胞死亡的组成部分，在从线虫到哺乳动物的进化过程中是保守的。
Table. Important events in the history of cell death Exp Oncol. 2012 Oct;34(3):146-52
| Year | Event | Classification | Refe-rence |
|---|---|---|---|
| 19th and early 20th C: DESCRIPTIONS OF CELL DEATH IN MOSTLY PHYSIOLOGICAL SITUATIONS | |||
| 1842 | Vogt describes resorption of notochord during amphibian metamorphosis, notes that it is physiological | Physiological; metamorphosis | |
| 1864 | Weismann: massive cell loss in insect metamorphosis; describes histolysis | Physiological; metamorphosis; description | |
| 1872 | Stieda: death of chondrocytes during ossification | Physiological; development | [33] |
| 1883 | Metchnikoff: loss of muscles during amphibian metamorphosis; activity of phagocytes | Physiological; metamorphosis | [3] |
| 1885 | Flemming: cell death in rabbit ovarian follicle (chromatolysis) is physiological | Physiological; development | |
| 1889 | Felix: denervated myocytes die | Pathological | [34] |
| 1890 | Arnheim: margination of chromatin | Description of apoptosis | [35] |
| 1906 | Collin: morphology of dying motor and sensory neurons in chicks | Physiological; development; | [36] |
| 1910 | von Recklinghausen: oncosis from ischemia or toxic agents | Non-physiological; necrosis | |
| 1914 | Gräper: Chromatolysis of all cells eliminated from organs; phagocytosis of neighboring cells | Physiological; apoptosis (not yet named) as general | [37] |
| 1934–1951: ACCEPTANCE OF CELL DEATH AS PHYSIOLOGICAL | |||
| 1934 | Hamburger: A factor that controls number of neurons | Physiological; development | |
| 1949 | Hamburger and Levi-Montalcini: emphasize death of neurons in CNS of embryos | Physiological; development | |
| 1951 | Gluecksmann reviews cell death in ontogenesis; a normal part of animal development | Physiological; development | |
| 1955–1973: PHYSIOLOGICAL MECHANISMS; DEFINITION OF PROGRAMMED CELL DEATH AND APOPTOSIS | |||
| 1955–1959 | De Duve and others recognize lysosomes and identify them as “suicide bags” | Physiological and pathological; mechanism | |
| 1961 | Bellairs: describes cell death in chick embryo | Physiological; description | [38] |
| 1964–1965 | Lockshin and Williams: Programmed cell death in insect metamorphosis | Physiological; mechanism; programmed cell death | |
| 1966 | Saunders: “The death clock is ticking;” cell death is a suicide | Physiological; development | |
| 1966 | Tata: Programmed cell death requires protein synthesis (confirmed 1969, 1971 in insect metamorphosis and thymocytes involution; later in neuronal death; otherwise not general) | Physiological; mechanism | |
| 1972 | Kerr, Wyllie, Currie: Apoptosis is general in morphology and among animals | Physiological; mechanism | |
| 1973 | Schweichel and Merker: classify types of death | Physiological | [39] |
| 1973–1992: MECHANISMS OF APOPTOSIS AND IDENTIFICATION OF GENETICS OF APOPTOSIS | |||
| 1976 | Skalka et al.: DNA fragmentation is typical of apoptosis | [40] | |
| 1976 | Sulston: nuc-1, a gene involved in cell death | Caenorhabditis | [41] |
| 1979 | Farber and Fisher: Free radicals can cause cell suicide | [42] | |
| 1982 | Horvitz et al.: Genetic paths of cell death | Caenorhabditis | [43] |
| 1983 | Hedgecock: mutants of cell death genes | Caenorhabditis | [44] |
| 1984 | Wyllie et al.: relate chromatin condensation to DNA fragmentation | [24] | |
| 1989–1992 | Identification of a cell surface receptor that can lead to cell death (Fas/APO-1/CD95) | ||
| 1986 | Ellis and Horvitz: ced-3 and ced-4, genes that can kill cells | Caenorhabditis | [45] |
| 1992 | Hengartner et al.: ced 9, a gene that can protect cells | Caenorhabditis | [46] |
| 1991–1994 | p53 can induce apoptosis | ||
| 1992 | Phosphatidylserine is a marker of apoptosis | [47] | |
| 1992–2000: Discovery of the genes of cell death | |||
| 1992 | Vaux et al.: Bcl-2 equivalent to ced-9 | [48] | |
| 1993 | Yuan et al.: CED-3 is related to ICE | ||
| 1993 | Baculovirus p35 can inhibit apoptosis | [49] | |
| 1994 | Hengartner and Horvitz: CED-9 is similar to Bcl-2 | [50] | |
| 1996 | Caspases defined | [50] | |
| 1996–1997 | Importance of release of cytochrome c for inducing apoptosis | [51] | |
| 1997 | Zou et al.: identify Apaf-1, similar to CED-4 | [52] | |
| 1998 | Del Peso et al.: EGL-1 inhibits CED-9 | [53] | |
| 2000 ff RECOGNITION OF DIFFERENT TYPES OF CELL DEATH | |||
| 1970’s | Most cell death is considered to be lysosomal; many papers | ||
| 1973 | Schweichel and Merker: classify types of death | Physiological | [43] |
| 1995 | Majno and Joris: Apoptosis, oncosis, necrosis | [54] | |
| Lockshin and Zakeri: Other types of cell death | [55] | ||
| 2005 | Broker et al.: cell death independent of caspases | [56] | |
| 2005 | Kroemer et al.: Classification of cell death | [57] | |
| 2008 | Degterev and Yuan: Cell death programs, including necroptosis | [58] | |
| Kroemer et al.: Role of autophagy | [59, 60] | ||
| 2008, 2012 | Klionsky et al.: Guidelines for evaluating autophagy | [61] | |
主要参考文献
Am J Pathol 1990; 136: 593–608
Nature 1993; 362: 847–9
Cell 1993; 75: 641–52
Programmed cell death and apoptosis — where it came from and where it is going: From Elie Metchnikoff to the control of caspases，Exp Oncol. 2012 Oct;34(3):146-52.
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