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Apoptosis

程序化细胞死亡或细胞凋亡是多细胞生物体的重要生理过程。细胞生长分裂和细胞死亡速度之间的均衡能够根据内部或外部参数动态调整细胞数量。例如,在脊椎动物的神经系统发育过程中,大约有一半的细胞在形成后不久就凋亡。在成年生物体中,这种平衡对于保持器官和组织的大小和功能等非常重要。此类平衡调节异常时常会导致癌症。细胞凋亡与细胞坏死不同。细胞坏死是指受影响的细胞破裂后,导致可能的损害性炎症反应,而细胞凋亡呈现为非常有组织的方式:细胞收缩凝集,内部结构分解,DNA 断裂为碎片。然后,周围的细胞或巨噬细胞迅速吞噬死亡细胞。

凋亡过程的核心是 caspase,一种半胱氨酸蛋白酶家族。它们生成为酶前体,由其他 caspase 转化为活性后裂解。当起始酶前体(例如酶前体 8、9、10)在接头蛋白的帮助下聚合时,触发此 caspase 级联反应,从而促进相互激活,减少低蛋白酶活性或酶前体构象变化的影响。已激活的 caspase 之后自由激活效应 caspase(例如 caspase 3、6 和 7),促进细胞凋亡。它们的效果进一步受 Bcl-2 家族蛋白质(例如 Bcl-2、Bcl-xL)和 IAP(细胞凋亡抑制剂,例如 BIRC1、XIAP)控制。

细胞凋亡过程遵循若干途径。外源性死亡受体途径通过配体诱导,这些配体结合到含有胞质死亡域的死亡受体蛋白家族(例如 FAS 和 TRAIL 受体)。内源性途径则是响应 DNA 损坏或线粒体应力,与癌症特别相关。除了这些标准细胞凋亡途径外,还有 caspase 非依赖性途径,由粒酶 B 和粒酶 A 等触发。这些 caspase 非依赖性途径已针对抑制 caspase 的病毒产生进化。

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Caspases

Mitochondrial

Inhibitor of Apoptosis

BIRC2 (Baculoviral IAP Repeat Containing 2):

NAIP (NLR Family, Apoptosis Inhibitory Protein):

XIAP (X-Linked Inhibitor of Apoptosis):

BIRC3 (Baculoviral IAP Repeat Containing 3):

EPR1 (Early-Phytochrome-Responsive1):

Receptor

FASL (Fas Ligand (TNF Superfamily, Member 6)):

TNFRSF1B (Tumor Necrosis Factor Receptor Superfamily, Member 1B):

CRADD (CASP2 and RIPK1 Domain Containing Adaptor with Death Domain):

CYLD (Cylindromatosis (Turban Tumor Syndrome)):

DR4 (Drought-Repressed 4 Protein):

FAS (TNF Receptor Superfamily, Member 6):

FADD (Fas (TNFRSF6)-Associated Via Death Domain):

RIPK1 (Receptor (TNFRSF)-Interacting serine-threonine Kinase 1):

TRAF1 (TNF Receptor-Associated Factor 1):

TRAF2 (TNF Receptor-Associated Factor 2):

TRAF3 (TNF Receptor-Associated Factor 3):

TRAF5 (TNF Receptor-Associated Factor 5):

TRADD (TNFRSF1A-Associated Via Death Domain):

TNFRSF10A (Tumor Necrosis Factor Receptor Superfamily, Member 10a):

TNFRSF10B (Tumor Necrosis Factor Receptor Superfamily, Member 10b):

TNFRSF12A (Tumor Necrosis Factor Receptor Superfamily, Member 12A):

TNFRSF1A (Tumor Necrosis Factor Receptor Superfamily, Member 1A):

Cytoplasmatic

Endoplasmatic Reticulum

TREX1 (three Prime Repair Exonuclease 1):

CAPN2 (Calpain 2, (M/II) Large Subunit):

NME1 (Non-Metastatic Cells 1, Protein (NM23A) Expressed in):

Granzyme

GZMA (Granzyme A (Granzyme 1, Cytotoxic T-Lymphocyte-Associated serine Esterase 3)):

Nuclear

DNA damage

ABL1 (C-Abl Oncogene 1, Non-Receptor tyrosine Kinase):

ATR (Ataxia Telangiectasia and Rad3 Related):

ATM (Ataxia Telangiectasia Mutated):

TRIM29 (Tripartite Motif Containing 29):

Apoptosis Markers

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