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| 组蛋白修饰在精子发生中的研究及对水生动物的启示 |
| Study of histone modification on spermatogenesis and its implications in aquatic animals |
| 投稿时间：2023-11-29 修订日期：2024-03-12 |
| DOI： |
| 中文关键词: 组蛋白 翻译后修饰 染色质重塑 精子发生 |
| 英文关键词: Histone post-translational modification chromatin remodeling, spermatogenesis |
| 基金项目:国家重点研发计划（2022YFD2400405）,国家自然科学基金项目（32072295） |
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| 中文摘要: |
| 两性生殖中，精子作为携带父本信息的载体，是物种延续的关键因素。精子发生经历精原细胞、初级和次级精母细胞、圆形精子、成熟精子阶段。在圆形精子形成成熟精子过程中，染色质进行重塑，细胞形态发生剧烈变化，其中组蛋白修饰和组蛋白变体在这些事件中发挥了重要作用，如甲基化主要与基因表达的激活或抑制有关；乙酰化激活转录活性并参与组蛋白沉积和DNA修复；磷酸化促进转录后修饰或参与DNA双链断裂修复；泛素化能够调节不同细胞途径中各式各样的蛋白质底物。组蛋白变体在调节染色体结构发挥重要功能，如组蛋白H1变体在分化过程中具有抑制转录的作用；组蛋白H2A和H2B变体在精子染色质包装过程中发挥特有功能；H3.3是H3最重要的变体，在细胞周期的各个时期都表达；组蛋白H4则是进化最慢的组蛋白之一，目前还没有发现其组蛋白变体。本文围绕组蛋白翻译后修饰，梳理了甲基化、乙酰化、磷酸化、泛素化等方面的最新进展和组蛋白变体在染色质重塑过程中的功能研究，随后针对各类组蛋白变体及其功能进行了总结，最后以半滑舌鳎为例简要介绍了这些研究对水生动物精子发生的启示。 |
| 英文摘要: |
| Epigenetics indicates the heritable changes that do not change the DNA sequence. Compared with genetic changes, epigenetic changes affect gene expression and protein products in cells, while the changes are reversible and dependent on the environment. There are three major types of epigenetic changes: DNA methylation, histone post-translational modification and non-coding RNA (ncRNA). In this paper, we will focus on the process of histone post-translational modification.
Histones have five main types: H1/H5, H2A, H2B, H3 and H4. The genes encoding histones do not contain introns and histones are one of the most conservative proteins in eukaryotes. Histone is the basic structural protein that makes up the eukaryotic chromosomes. In general, two molecules of H2A, H2B, H3 and H4 form a histone octamer, which combines with DNA to form a structural unit called nucleosome. This nucleosome appears in every 200 base pairs, which is connected by H1 histones to form chromatin.
Histone modification refers to addition of a functional group to the amino acids (most commonly lysine) in the histone tail. The histone modification can control gene expression through chromatin condensation and depolymerization, and can also provide binding sites for various proteins. Histone modifications reported in animals include methylation, acetylation, phosphorylation, ubiquitin, SUMOylation, ADP ribosylation, short-chain lysine acylation and so on.
A large number of studies have shown that chromatin remodeling is a key step in spermatogenesis, involving the transformation of histone to protamine. In short, the process of protamine replacing histone requires (I) histone post-translational modification (PTM) to promote the opening of histone-based chromatin structures, especially histone hyperacetylation and incorporation into histone variants, (II) Bromine domain proteins binding to acetyl residues and remodeling the chromatin, (III)the formation and repair of DNA strand breaks in chromatin remodeling, and (Iv) incorporation of protamine. Here we will focus on process (I).
In bisexual reproduction, sperm, as a carrier of paternal information, is a key factor in the continuation of species. Spermatogenesis includes various stages, spermatogonia, primary and secondary spermatocytes, round sperm and mature sperm. In the process of round sperm transforming to mature sperm, chromatin is remodeling occurs and cell morphology experiences dramatic changes, in which histone post-translational modification and histone variants play an important role in these events. Histone post-translational modification patterns affect gene expression at a wide range, such as methylation is mainly related to the activation or inhibition of gene expression, acetylation activates transcriptional activity and participates in histone deposition and DNA repair, phosphorylation promotes post-transcriptional modification or participates in DNA double strand break repair, ubiquitin can regulate a variety of protein substrates in different cellular pathways. Histone variants have special functions in regulating chromosome structure, such as histone H1 variants inhibit transcription during differentiation, histone H2A and H2B variants play a unique role in sperm chromatin packaging, H3.3 is the most important variant of H3, which is expressed in all stages of the cell cycle and can participate in chromosome formation outside the s phase, Histone H4 as one of the slowest evolving proteins, no histone variant has been found so far. Focusing on the post-translational modification of histone, this paper reviews the latest progress in methylation, acetylation, phosphorylation, ubiquitination. Subsequently, the histone variants and their function in the process of chromatin remodeling are summarized. Finally, taking Cynoglossus semilaevis as an example, this paper briefly introduces the implications of these studies on spermatogenesis in aquatic animals. Elucidating the effect of post-translational modifications in spermatogenesis will help to explore the regulatory mechanism of the specific sperm (W type) absence, which would not only expand the fundamental theory of reproduction biology, but also provide new solutions for the cultivation of monosex fry in aquaculture. |
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