昆虫学报 ›› 2023, Vol. 66 ›› Issue (2): 245-254.doi: 10.16380/j.kcxb.2023.02.014

• 综 述 • 上一篇    下一篇

昆虫性别决定级联的研究进展

刘雅婷1, 王文璐2, 陈宗泽1, 谢文2,*, 张友军2,*   

  1. (1. 佛山科学技术学院食品科学与工程学院, 佛山 528000; 2. 中国农业科学院蔬菜花卉研究所, 北京 100081)
  • 出版日期:2023-02-20 发布日期:2023-04-07

Research progress on sex determination cascade in insects

LIU Ya-Ting1, WANG Wen-Lu2, CHEN Zong-Ze1, XIE Wen2,*, ZHANG You-Jun2,*   

  1. (1. School of Food Science and Engineering, Foshan University, Foshan 528000, China; 2. Institute of Vegetables and Flowers, Chinese Academy of Agricultural Sciences, Beijing 100081, China)
  • Online:2023-02-20 Published:2023-04-07

摘要:  性别决定是发育和进化生物学研究的一个重大问题。已知大多数昆虫的性别决定级联为:初级信号因子→性别决定关键基因→双性基因→性别分化基因。尽管遵循这样的模式,但不同昆虫的性别决定基因和调控机制各不相同,特别是性别决定初级信号因子存在较大分歧。自黑腹果蝇Drosophila melanogaster的初级信号被发现以来,人们陆续确定了蚊子、蜜蜂、丽蝇蛹集金小蜂Nasonia vitripennis、家蚕Bombyx mori等模式昆虫的初级信号。初级信号的种类复杂多样,包括性染色体的剂量、雄性化因子(male-determining factors, M factors)、等位基因的杂合度、母代印记等,这在一定程度上增加了非模式昆虫的研究难度。尽管如此,昆虫性别决定级联的下游调控机制仍相对保守,特别是transformer(tra)+transformer2(tra2)→doublesex(dsx)/fruitless(fru)的调控模式在大多数昆虫中存在共性。tra通过感知初级信号而发生选择性可变剪接,并在tra2的帮助下实现其对自身及下游dsxfru的剪接调控,从而维持性别发育。dsx作为级联末端的“双开关”,是昆虫性别决定级联中最保守的基因。它在调控两性发育、求偶行为、外生殖器及性二态特征的形成方面均表现出高度的保守性。fru作为果蝇性行为的开关基因,几乎参与调控果蝇所有雄性性行为。它的功能在橘小实蝇Bactrocera dorsalis、埃及伊蚊Aedes aegypti、家蚕等多种昆虫中得到验证,成为理解昆虫复杂性行为的典型基因。了解昆虫性别决定级联,理清各性别决定基因的功能及其相互作用关系,是阐明性别决定机制的根本,为揭示昆虫性别决定的一般规律、推动昆虫性别决定上游调控机制的基础研究以及实现昆虫的性别遗传操控提供理论基础。

关键词: 昆虫, 性别决定级联, 初级信号, 关键基因, 可变剪接

Abstract:  Sex determination is an outstanding question in developmental and evolutionary biology. The sex determination cascade in most of known insects is: primary signaling element→key gene of sex determination→double switch gene→sex differentiation gene. In spite of this pattern, different insects have different sex determination genes and regulatory mechanisms, especially the primary signaling element of sex determination. Since the primary signal of Drosophila melanogaster was discovered, the primary signal of mosquito, bee, wasp (Nasonia vitripennis), silkworm (Bombyx mori) and other model insects has been determined successively. There are many kinds of primary signals, including the dose of sex chromosomes, male-determining factors (M factors), heterozygosity of alleles and maternal imprinting, which make it more difficult to study non-model insects to some extent. Even so, the downstream regulatory mechanism is relatively conserved. In particular, the transformer (tra)+transformer2 (tra2)→doublesex (dsx)/fruitless (fru) pathway is fairly common in most insects. Tra produces alternative splicing by sensing primary signals, and with the help of tra2, tra regulates the splicing of itself and the downstream dsx and fru, and maintains gender development. Acting as the terminal ‘double-switch’, dsx is the most conserved gene in the insect sex determination cascade. dsx is highly conserved in regulating bisexual development, courtship behavior, genitalia and formation of sexual dimorphism. As a switch gene of Drosophila sexual behavior, fru is involved in regulating almost all male sexual behaviors of Drosophila. Its function has been verified in a variety of insects such as Bactrocera dorsalis, Aedes aegypti and B. mori, and it has become a typical gene for understanding the complex sexual behavior of insects. Understanding the sex determination cascade in insects, and clarifying the function and interaction of each sex determination gene are essential to elucidate the mechanism of sex determination. It provides a theoretical basis for revealing the general law of insect sex determination, and promoting the basic research on the upstream regulatory mechanism of insect sex determination, and realizing the genetic manipulation of insect sex determination.

Key words: Insect, sex determination cascade, primary signal, key gene; alternative splicing