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浙江大学学报（农业与生命科学版）
Journal of Zhejiang University (Agriculture and Life Sciences)  2023, Vol. 49 Issue (2): 213-228    DOI: 10.3785/j.issn.1008-9209.2022.03.101
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Review on programmed cell death and vertebrate embryonic development
Jingyun LUAN(),Pengfei XU()
College of Medicine, Zhejiang University, Hangzhou 310058, Zhejiang, China
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Abstract  

Vertebrate embryonic development is a complex process in which cells self-organize into a completely formed organism via cell behaviors such as division, proliferation, differentiation, migration, and programmed cell death (PCD). Among these cell behaviors, PCD exists in all stages of embryonic development, playing important roles in organogenesis, morphogenesis, and maintenance of tissue homeostasis. In this review, we summarized recent research progress on the regulation of vertebrate embryonic development by PCD, including the biological function and regulatory mechanism of PCD and clearance of dead cells during early embryonic development, focusing on the close link between PCD and embryonic developmental processes. We hope this review will help build a more comprehensive understanding of the roles of PCD in vertebrate embryonic development, and provide insights into how PCD is manipulated to improve the quality of embryonic development in the future.

Key words： programmed cell death      vertebrate      embryonic development     
Received: 10 March 2022      Published: 27 April 2023
CLC:  Q593.4  
Corresponding Authors: Pengfei XU     E-mail: 21918064@zju.edu.cn;pengfei_xu@zju.edu.cn
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Cite this article:

Jingyun LUAN,Pengfei XU. Review on programmed cell death and vertebrate embryonic development. Journal of Zhejiang University (Agriculture and Life Sciences), 2023, 49(2): 213-228.

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https://www.zjujournals.com/agr/10.3785/j.issn.1008-9209.2022.03.101     OR     https://www.zjujournals.com/agr/Y2023/V49/I2/213


细胞程序性死亡与脊椎动物胚胎发育综述

脊椎动物胚胎发育是细胞通过分裂、增殖、分化、迁移及细胞程序性死亡等一系列行为进行自我组织的复杂过程。其中，细胞程序性死亡存在于胚胎发育的各个阶段，对胚胎的发育具有重要调控作用，是器官生成、形态建立和组织稳态维持等发育相关事件所必需的。本文对近年来细胞程序性死亡调控脊椎动物胚胎发育过程的相关研究进展进行了综述，包括早期胚胎发育过程中细胞程序性死亡的生物学功能、调节机制及死亡细胞的清除方式等，重点关注了细胞程序性死亡与脊椎动物胚胎发育过程之间的密切联系。本综述将为更全面地了解细胞程序性死亡在脊椎动物胚胎发育过程中的作用提供借鉴与帮助，并有望为将来通过人为调控细胞程序性死亡来提高胚胎发育质量提供一定的思路。


关键词： 细胞程序性死亡,  脊椎动物,  胚胎发育 

细胞死亡类型

Cell death type

形态学特征

Morphological characteristic

生化特征

Biochemical characteristic

细胞凋亡 Apoptosis

核碎裂，质膜起泡，染色质凝聚，细胞固缩，凋

亡小体形成

半胱天冬酶被激活，磷脂酰丝氨酸暴露，线粒体跨膜电位（ΔΨm）耗散
细胞自噬 Autophagy双膜自噬泡积累，细胞质空泡化，无染色质凝聚脂化微管相关蛋白轻链3-Ⅰ（LC3-Ⅰ）向LC3-Ⅱ转化，底物降解
细胞坏死 Necrosis质膜破裂，细胞及细胞器肿胀，染色质浓缩腺苷三磷酸（ATP）耗竭，钙蛋白酶和组织蛋白酶参与的蛋白水解，损伤相关分子模式（DAMP）分子释放
铁死亡 Ferroptosis

线粒体变小且密度增加，嵴减少或消失，线粒

体外膜破裂

铁离子及活性氧积累，胱氨酸/谷氨酸反向转运体（system Xc－）被激活，谷胱甘肽被消耗，脂质过氧化
细胞焦亡 Pyroptosis质膜破裂，细胞轻度肿胀caspase及消皮素D被激活，大量促炎症因子释放
NETosis核膜分解产生小囊泡，染色质去浓缩中性粒细胞弹性蛋白酶及髓过氧化物酶释放，PAD4被激活
铜死亡 Cuprotosis脂酰化蛋白质聚集，铁硫簇蛋白减少铜离子与脂酰化修饰的蛋白质结合
Table 1 Cell death types and their characteristics
Fig. 1 Biological function of programmed cell death during mouse embryonic development
Fig. 2 Programmed cell death regulates proper tissue and organ generationA. Programmed cell death regulates the establishment of the correct morphogen concentration gradient; B. Programmed cell death is involved in the production of tissues and organs; C. Programmed cell death is involved in the elimination of temporary structures or organs; D. Programmed cell death controls the total number of cells in the body.
Fig. 3 Elimination ways of early dead cellA. Epithelial cell extrusion eliminates dead cells; B1-B2. Epithelial and neural crest cells recognize and phagocytose dead cells (B1. Epithelial cells induce rapid dispersal and uptake of apoptotic targets through phagocytic cups and ‘epithelial arms’, and the mechanical load-sharing mechanism enables the long-range cooperative uptake of apoptotic cells by multiple epithelial cells load-sharing; B2. Neural crest cells respond rapidly to dying cells around the neural tube, and then migrate from its segmentally-restricted paths towards dead cells and phagocytose debris).
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