黑腹果蝇转座子的特性、调控及其在基因组进化中的作用

doi:10.12300/j.issn.1674-5817.2025.112

实验动物与比较医学 ›› 2025, Vol. 45 ›› Issue (6): 676-687.DOI: 10.12300/j.issn.1674-5817.2025.112

• 无脊椎实验动物：果蝇 • 上一篇下一篇

黑腹果蝇转座子的特性、调控及其在基因组进化中的作用

王也, 王露()()

核糖核酸功能与应用全国重点实验室, 中国科学院分子细胞科学卓越创新中心/生物化学与细胞生物学研究所, 中国科学院大学, 上海 200031

收稿日期:2025-07-07 修回日期:2025-08-12 出版日期:2025-12-25 发布日期:2025-12-19
通讯作者: 王露（1989—），男，博士，研究员，研究方向：转座子的调控机制和功能。E-mail：lu.wang@sibcb.ac.cn。ORCID：0000-0003-2356-6245
作者简介:王也（2000—），女，硕士研究生，研究方向：转座子转录调控机制研究。E-mail：wangye2022@sibcb.ac.cn
基金资助:
国家自然科学基金面上项目“多梳蛋白Crm抑制逆转座子的分子机制探究”(32270600)

Drosophila melanogaster Transposons: Characterization, Regulation, and Their Role in Genome Evolution

WANG Ye, WANG Lu()()

Key Laboratory of RNA Innovation, Science and Engineering, Center for Excellence in Molecular Cell Science/Shanghai Institute of Biochemistry and Cell Biology, University of Chinese Academy of Sciences, Shanghai 200031, China

Received:2025-07-07 Revised:2025-08-12 Published:2025-12-25 Online:2025-12-19
Contact: WANG Lu (ORCID: 0000-0003-2356-6245), E-mail: lu.wang@sibcb.ac.cn

摘要/Abstract

摘要：

转座子（transposable elements，TEs）是基因组中可移动的DNA序列，在物种演化、基因组稳定性及基因调控中扮演关键角色。黑腹果蝇（Drosophila melanogaster）作为经典模式动物，其基因组中TEs占比约20%，是研究TEs的生物学特性、宿主防御机制及功能演化的理想模型，也为理解高等生物乃至人类TEs相关疾病的机制提供了重要范式。本文系统阐述了黑腹果蝇中TEs的分类、分布特征及其与宿主基因组的动态互作，重点探讨了以PIWI相互作用RNA（PIWI-interacting RNA，piRNA）通路为核心的宿主防御系统。接着，本文详细解析了黑腹果蝇中关键TEs家族（如Gypsy、Copia、P-element、I-element）的生物学特性及其在基因组进化中的双重作用：一方面，TEs插入可引发基因组不稳定、杂种不育及衰老表型，为研究相关人类疾病（如神经退行性疾病、基因组不稳定综合征等）提供了模型基础；另一方面，其序列可被宿主驯化（co-option）为新型调控元件或功能基因，驱动适应性创新。最后，本文还展望了TEs未来研究方向，包括环境应激对TEs活性的调控、piRNA通路与其他小RNA系统的交互，以及TEs在衰老和神经退行性疾病发生发展中的调控效应。黑腹果蝇TEs的研究不仅深化了人们对TEs生物学的理解，其揭示的保守机制和原理，还为利用实验动物模型研究人类疾病、开发基因治疗和基因编辑技术提供了关键理论基础和重要启示。

关键词: 黑腹果蝇, 转座子, piRNA, 功能演化

Abstract:

Transposable elements (TEs) are mobile DNA sequences in genomes that play key roles in species evolution, genome stability, and gene regulation. Drosophila melanogaster, as a classic model animal with TEs accounting for approximately 20% of its genome, is an ideal model for studying biological characteristics, host defense mechanisms, and functional evolution of TEs, and also provides an important paradigm for understanding mechanisms of TE-related diseases in higher organisms and even humans. This review systematically elucidates the classification and distribution characteristics of TEs in D. melanogaster and their dynamic interactions with host genome, focusing on the host defense system centered on PIWI-interacting RNA (piRNA) pathway. Then, the biological characteristics of key TE families (such as Gypsy, Copia, P-element, and I-element) in D. melanogaster and their dual roles in genomic evolution are analyzed in detail. On the one hand, TE insertions can cause genomic instability, heterozygous sterility, and aging phenotypes, providing a model basis for studying related human diseases (e.g., neurodegenerative diseases, genomic instability syndromes, etc.). On the other hand, their sequences can be co-opted by the host to create novel regulatory elements or functional genes, thereby driving adaptive innovation. Finally, the future research directions of TEs are proposed, including regulation of TE activity by environmental stress, interaction between piRNA pathways and other small RNA systems, as well as regulatory effects of TEs in the occurrence and development of aging and neurodegenerative diseases. The research on TEs in D. melanogaster not only deepens understanding of TE biology, but also provides a key theoretical basis and important inspiration for studying human diseases using experimental animal models, as well as for developing gene therapy and gene editing technologies.

Key words: Drosophila melanogaster, Transposable elements, piRNA, Functional evolution

中图分类号:

Q95-33

王也,王露. 黑腹果蝇转座子的特性、调控及其在基因组进化中的作用[J]. 实验动物与比较医学, 2025, 45(6): 676-687. DOI: 10.12300/j.issn.1674-5817.2025.112.

WANG Ye,WANG Lu. Drosophila melanogaster Transposons: Characterization, Regulation, and Their Role in Genome Evolution[J]. Laboratory Animal and Comparative Medicine, 2025, 45(6): 676-687. DOI: 10.12300/j.issn.1674-5817.2025.112.

图/表 3

图1 果蝇转座子的结构与分类示例注：黑色方框表示开放阅读框；彩色区域表示定义的蛋白质结构域；折线段表示内含子；三角形表示重复序列；颜色相同（灰色除外）的结构域表示有共同的祖先；AP，无嘌呤/无嘧啶；ATPase，腺苷三磷酸酶；Cys，半胱氨酸；DJR MCP，双果冻卷主要衣壳蛋白；EN，核酸内切酶；IN，整合酶；ORF，开放阅读框；pPolB，蛋白引发型DNA聚合酶B；PR，pol编码蛋白酶；RH，核糖核酸酶H结构域；RT，逆转录酶；SJR mCP，单果冻卷结构小衣壳蛋白。

Figure 1 Examples of the structure and classification of Drosophila transposable elementsNote： Black boxes indicate open reading frames； Colored regions indicate defined protein structural domains； Folded segments indicate introns； Triangles indicate repeated sequences； Structural domains of the same color （excluding gray） share a common ancestor； AP， apurinic/apyrimidinic； ATPase， adenosine triphosphatase； Cys， cysteine； DJR MCP， double jelly-roll major capsid protein； EN， endonuclease； IN， integrase； ORF， open reading frame； pPolB， protein-primed type B DNA polymerase； PR， pol-encoded protease； RH， Ribonuclease H domain； RT， reverse transcriptase； SJR mCP， single jelly-roll minor capsid protein.

图2 piRNA的生物发生示意图注：2'-O-me，2'-O-甲基化；Me，甲基；piRNA，PIWI相互作用RNA；TE，转座子；Ago3，Argonaute 3；Armi，Armitage；Aub，Aubergine；Hsp90，热休克蛋白90；Mino，Minotaur；Vret，纺锤蛋白；Zuc，Zucchini；Hen1，Hen1甲基转移酶；Vasa，Vasa DEAD-box RNA解旋酶；Gasz，种系相关支架锌指蛋白。

Figure 2 Schematic diagram of piRNA biogenesisNote： 2'-O-me， 2'-O-methylation； Me， methyl group； piRNA， PIWI-interacting RNA； TE， transposable element； Ago3， Argonaute 3； Armi， Armitage； Aub， Aubergine； Hsp90， Heat shock protein 90； Mino， Minotaur； Vret， Vreteno； Zuc， Zucchini； Hen1， Hen1 methyl-transferase； Vasa， Vasa DEAD-box RNA helicase； Gasz， Germline-associated scaffolding zinc finger.

表1 果蝇中TEs宿主驯化的主要类型、机制与功能效应实例

Table 1 Examples of major types, mechanisms and functional effects of TEs host co-option in Drosophila

驯化类型 Type of co-option	驯化机制/来源 Mechanism/Source of co-option	功能影响 Functional impact	代表性案例与功能 Representative case & function
调控元件驯化 Regulatory element co-option	TEs调控序列（如启动子、增强子、绝缘子）	改变邻近/宿主基因的表达模式、组织特异性、发育时序或环境响应性	果蝇体色演化：TEs（尤其LTR型）插入影响色素相关基因的调控，驱动体色模式快速变化^[18,27]
编码序列驯化（完全蛋白） Protein-coding sequence co-option (full-length)	TEs完整或近乎完整的编码序列	产生具有细胞功能的全新结构蛋白或酶	PIF-like基因：果蝇属中多次独立驯化自PIF转座酶，形成新的功能基因家族^[20]
编码序列驯化（功能域/模块） Protein-coding sequence co-option (functional domain/module)	TEs编码蛋白的特定结构域	赋予宿主蛋白新功能或参与新细胞过程	Arc/Arc1蛋白（gag域）：来源于逆转录TEs（如Gypsy、Copia）GAG蛋白，形成衣壳样结构，包装自身mRNA，介导神经元-肌肉或神经元间RNA运输，对突触可塑性和记忆维持至关重要^[67-68]
整体元件利用 Holistic element utilization	特定TEs家族的活性或产物	在特定生理或发育过程中被宿主利用	mdg4（Gypsy家族）：在果蝇蛹期变态窗口期激活，通过STING–Relish通路诱导全身性抗病毒免疫状态^[69]

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黑腹果蝇转座子的特性、调控及其在基因组进化中的作用

Drosophila melanogaster Transposons: Characterization, Regulation, and Their Role in Genome Evolution

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