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Silencing transformer and transformer-2 in Zeugodacus cucurbitae causes defective sex determination with inviability of most pseudomales

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Journal of Integrative Agriculture 2024年第3期23卷 938-947页

作者： Qin Ma Zizhen Fan Ping Wang Siya Ma Jian Wen Fengqin Cao Xianwu Lin Rihui Yan Key Laboratory of Green Prevention and Control of Tropical Plant Diseases and Pests of Ministry of Education School of Plant ProtectionHainan UniversityHaikou 570228China School of Life Sciences HainanUniversityHaikou 570228China Hainan Yazhou Bay Seed Lab Sanya 572000China

transformer is a switch gene for sex determination in many insects, which cooperates with transformer-2 that is expressed in both sexes to regulate female differentiation, particularly in dipterans. Zeugodacus cucurbitae(Coquillett) is a very destructive pest worldwide, however, its sex determination pathway remains largely uncharacterized. Here, we show that the female sex ratio is sharply reduced with knockdown of either transformer or transformer-2 by RNA interference in early embryos of Z. cucurbitae. Most of the males grown from the embryos with transient transformer and transformer-2 suppression mated with wild-type females and produced mixed sex progeny, with one exception that produced only female progeny, and all of the few remaining males failed to mate with wild-type females and produced no progeny. The exceptional male and those males with mating failure were XX pseudomales as determined by the detection of Y chromosome-linked Maleness-on-the-Y, indicating that most XX pseudomales are not viable. The phenotypes of transformer and transformer-2 suggest that they play a key role in regulating sex determination and are required for female sexual development of Z. cucurbitae. Our results will be beneficial to the understanding of sex determination in Z. cucurbitae and can facilitate the development of genetic sexing strains for its biological control.

关键词： Zeugodacus cucurbitae transformer transformer-2 sex determination RNA interference biological control

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sex determination in Homo sapiens as a Multi-Step Process: Potential for Development of Variants and sex Differences in Disease Risk

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Journal of Biomedical Science and Engineering 2024年第1期17卷 13-34页

作者： David A. Hart Department of Surgery Faculty of Kinesiology McCaig Institute for Bone & Joint Health University of Calgary Cal-gary Canada

Reproduction via cis-binary mechanisms appears to have evolved fairly early in the evolution of complex organisms, and a system committed to prior to evolution of humans. While the evolution of a chromosomal-specific approach has been a successful strategy for survival of a large variety of species including humans, the fidelity of sex determination leading to 100% cis-binary outcomes is not achieved in many species, with evidence for homosexual or bisexual behaviour evident in more than 1500 species. Thus, such outcomes indicates that sex determination is a multi-step process and not a single event, and as such, could lead to the appearance of variants during the process which developed much earlier than humans. Variants could arise either due to intrinsic variation in the steps of determination, or also be influenced by environmental factors of a biological or psychological nature. In contrast to homosexual variants which do not require interventions such as hormone therapy or surgery, expression of gender dysphoria, is more based in psychology, but also has biological underpinnings and can be influenced by such hormonal interventions and surgery. While the numbers of those with gender dysphoria is small (~0.6% - 1.0% of population), the attention given to this issue raises the possibility of biological and psychological environmental factors impacting the emergence of some of those expressing gender dysphoria. Furthermore, transitioning from male-to-female or female-to-male can have consequences regarding disease risks latter in life, including the appearance of autoimmune diseases. This review will attempt to review some of the evidence regarding sex determination, discuss why the system has potentially not been improved upon during evolution, how a potential role for sex chromosome function on neurodevelopment may be central to variation in humans, and how commitment to the current strategy is likely integrated into other sex-related events such as puberty, pregnancy, and menopause to ensure species survival

关键词： sex determination Variant Development Fidelity of sex determination Biological sex determination Species Variation

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sex determination mechanisms and sex control approaches in aquaculture animals

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Science China(Life Sciences) 2022年第6期65卷 1091-1122页

作者： Xi-Yin Li Jie Mei Chu-Tian Ge Xiao-Li Liu Jian-Fang Gui State Key Laboratory of Freshwater Ecology and Biotechnology Hubei Hongshan LaboratoryThe Innovative Academy of Seed DesignInstitute of HydrobiologyChinese Academy of SciencesUniversity of Chinese Academy of SciencesWuhan 430072China College of Fisheries Hubei Hongshan LaboratoryHuazhong Agricultural UniversityWuhan 430070China College of Biological and Environmental Sciences Zhejiang Wanli UniversityNingbo 315100China Key Laboratory of Tropical&Subtropical Fishery Resource Application&Cultivation of Ministry of Agriculture and Rural Affairs Pearl River Fisheries Research InstituteChinese Academy of Fishery SciencesGuangzhou 510380China

Aquaculture is one of the most efficient modes of animal protein production and plays an important role in global food security.Aquaculture animals exhibit extraordinarily diverse sexual phenotypes and underlying mechanisms,providing an ideal system to perform sex determination research,one of the important areas in life science.Moreover,sex is also one of the most valuable traits because sexual dimorphism in growth,size,and other economic characteristics commonly exist in aquaculture animals.Here,we synthesize current knowledge of sex determination mechanisms,sex chromosome evolution,reproduction strategies,and sexual dimorphism,and also review several approaches for sex control in aquaculture animals,including artificial gynogenesis,application of sex-specific or sex chromosome-linked markers,artificial sex reversal,as well as gene editing.We anticipate that better understanding of sex determination mechanisms and innovation of sex control approaches will facilitate sustainable development of aquaculture.

关键词： sex determination sex control approach sexual dimorphism sex chromosome sex change reproduction strategy sex reversal

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Early embryonic transcriptomes of Zeugodacus tau provide insight into sex determination and differentiation genes

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Insect Science 2022年第3期29卷 915-931页

作者： Peipei Liu Wenping Zheng Jiao Qiao Ziniu Li Zhurong Deng Yimin Yuan Hongyu Zhang Key Laboratory of Horticultural Plant Biology(MOE) Hubei Hongshan LaboratoryChina-Australia Joint Research Center for Horticultural and Urban PestsInstitute of Urban and Horticultural EntomologyCollege of Plant Science and TechnologyHuazhong Agricultural UniversityWuhanChina College of Environmental Design Wuhan Institute of Design and SciencesWuhanChina

Zeugodacus tau(Walker)is an invasive pest.The sterile insect technique is an environment-friendly method for pest control.Understanding the mechanism of sex determination will contribute to improving efficiency of this technique.In this study,we identified the transformer(tra)gene in Z.tau.One female-specific and two male-specific isoforms of tra were found in Z.tau,and the male-specific splicing pattern of tra was found to occur 5 h after egg laying.We performed transcriptome sequencing at 1 h(E1),5 h(E5),and 9 h(E9)after egg laying and obtained high-quality transcriptome libraries of early embryo development.We identified 13297 and 11713 differentially expressed genes(DEGs)from E5 versus E1 and E9 versus E1 comparisons,respectively.To explore the potential functions of the DEGs during embryonic development,Gene Ontology,Clusters of Orthologous Groups of proteins,and Kyoto Encyclopedia of Genes and Genomes analyses were performed.Twenty-six genes potentially involved in sex determination or differentiation,including Maleness-on-the-Y(MoY),were identified in Z.tau.To verify the transcriptome results,15 genes were selected for quantitative real-time PCR validation.The results were consistent with the transcriptome sequencing results.Moreover,U2 small nuclear riboprotein auxiliary factor(U2AF-50),female lethal d(fl(2)d),and virilizer(vir)were highly expressed at E5,indicating that they may be related to the sex-specific splicing of tra.Further functional analysis is needed to confirm this speculation.Our data provide an insight into the mechanism underlying sex determination and differentiation in tephritid species.

关键词： RNA-Seq sex determination sex differentiation transformer Zeugodacus tau

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Chromosome-level Genomes Reveal the Genetic Basis of Descending Dysploidy and sex determination in Morus Plants

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Genomics, Proteomics & Bioinformatics 2022年第6期20卷 1119-1137页

作者： Zhongqiang Xia Xuelei Dai Wei Fan Changying Liu Meirong Zhang Peipei Bian Yuping Zhou Liang Li Baozhong Zhu Shuman Liu Zhengang Li Xiling Wang Maode Yu Zhonghuai Xiang Yu Jiang Aichun Zhao State Key Laboratory of Silkworm Genome Biology Institute of Sericulture and Systems BiologySouthwest UniversityChongqing 400716China Key Laboratory of Animal Genetics Breeding and Reproduction of Shaanxi ProvinceCollege of Animal Science and TechnologyNorthwest A&F UniversityYangling 712100China Key Laboratory of Coarse Cereal Processing Ministry of Agriculture and Rural AffairsChengdu UniversityChengdu 610106China The Sericultural and Apicultural Research Institute Yunnan Academy of Agricultural SciencesMengzi 661100China College of Sericulture Textile and Biomass SciencesSouthwest UniversityChongqing 400716China

Multiple plant lineages have independently evolved sex chromosomes and variable karyotypes to maintain their sessile lifestyles through constant biological innovation.Morus notabilis,a dioecious mulberry species,has the fewest chromosomes among Morus spp.,but the genetic basis of sex determination and karyotype evolution in this species has not been identified.In this study,three high-quality genome assemblies were generated for Morus spp.[including dioecious M.notabilis(male and female)and Morus yunnanensis(female)]with genome sizes of 301–329 Mb and were grouped into six pseudochromosomes.Using a combination of genomic approaches,we found that the putative ancestral karyotype of Morus species was close to 14 protochromosomes,and that several chromosome fusion events resulted in descending dysploidy(2n=2x=12).We also characterized a~6.2-Mb sex-determining region on chromosome 3.Four potential male-specific genes,a partially duplicated DNA helicase gene(named MSDH)and three Ty3_Gypsy long terminal repeat retrotransposons(named MSTG1/2/3),were identified in the Y-linked area and considered to be strong candidate genes for sex determination or differentiation.Population genomic analysis showed that Guangdong accessions in China were genetically similar to Japanese accessions of mulberry.In addition,genomic areas containing selective sweeps that distinguish domesticated mulberry from wild populations in terms of flowering and disease resistance were identified.Our study provides an important genetic resource for sex identification research and molecular breeding in mulberry.

关键词： Mulberry Karyotype evolution Dioecy sex determination Population genomics

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Evolution of sex determination in crustaceans

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Marine Life Science & Technology 2023年第1期5卷 1-11页

作者： Zhiqiang Ye Trent Bishop Yaohai Wang Ryan Shahriari Michael Lynch Center for Mechanisms of Evolution Arizona State UniversityTempeAZ 85287USA Institute of Evolution and Marine Biodiversity KLMMEOcean University of ChinaQingdao 266003China

sex determination(SD)involves mechanisms that determine whether an individual will develop into a male,female,or in rare cases,hermaphrodite.Crustaceans harbor extremely diverse SD systems,including hermaphroditism,environmental sex determination(ESD),genetic sex determination(GSD),and cytoplasmic sex determination(e.g.,Wolbachia controlled SD systems).Such diversity lays the groundwork for researching the evolution of SD in crustaceans,i.e.,transitions among different SD systems.However,most previous research has focused on understanding the mechanism of SD within a sin-gle lineage or species,overlooking the transition across different SD systems.To help bridge this gap,we summarize the understanding of SD in various clades of crustaceans,and discuss how different SD systems might evolve from one another.Furthermore,we review the genetic basis for transitions between different SD systems(i.e.,Dmrt genes)and propose the microcrustacean Daphnia(clade Branchiopoda)as a model to study the transition from ESD to GSD.

关键词： Crustacean sex determination sexual differentiation Dmrt Daphnia

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Absence of complementary sex determination in two Leptopilina species(Figitidae,Hymenoptera)and a reconsideration of its incompatibility with endosymbiont-induced thelytoky

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Insect Science 2022年第3期29卷 900-914页

作者： Fangying Chen Martijn Schenkel Elzemiek Geuverink Louis van de Zande Leo W.Beukeboom Groningen Institute for Evolutionary Life Sciences University of GroningenP.O.Box 11103 Groningen9700 CCthe Netherlands

Complementary sex determination(CSD)is a widespread sex determination mechanism in haplodiploid Hymenoptera.Under CSD,sex is determined by the allelic state of one or multiple CSD loci.Heterozygosity at one or more loci leads to female development,whereas hemizygosity of haploid eggs and homozygosity of diploid eggs results in male development.sexual(arrhenotokous)reproduction normally yields haploid male and diploid female offspring.Under asexual reproduction(thelytoky),diploidized unfertilized eggs develop into females.Thelytoky is often induced by bacterial endosymbionts that achieve egg diploidization by gamete duplication.As gamete duplication leads to complete homozygosity,endosymbiont-induced thelytokous reproduction is presumed to be incompatible with CSD,which relies on heterozygosity for female development.Previously,we excluded CSD in four Asobara(Braconidae)species and proposed a two-step mechanism for Wolbachia-induced thelytoky in Asobara japonica.Here,we conclusively reject CSD in two cynipid wasp species,Leptopilina heterotoma and Leptopilina clavipes.We further show that thelytoky in L.clavipes depends on Wolbachia titer but that diploidization and feminization steps cannot be separated,unlike in A.japonica.We discuss what these results reveal about the sex determination mechanism of L.clavipes and the presumed incompatibility between CSD and endosymbiont-induced thelytoky in the Hymenoptera.

关键词： arrhenotoky haplodiploidy inbreeding sex determination thelytoky Wolbachia bacteria

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Maleness-on-the-Y(MoY) orthologue is a key regulator of male sex determination in Zeugodacus cucurbitae(Diptera:Tephritidae)

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Journal of Integrative Agriculture 2023年第2期22卷 505-513页

作者： FAN Zi-zhen MA Qin MA Si-ya CAO Feng-qin YAN Ri-hui LIN Xian-wu Key Laboratory of Green Prevention and Control of Tropical Plant Diseases and Pests of Ministry of Education Hainan UniversityHaikou 570228P.R.China School of Plant Protection Hainan UniversityHaikou 570228P.R.China School of Life Sciences Hainan UniversityHaikou 570228P.R.China Hainan Yazhou Bay Seed Lab Sanya 572000P.R.China

The initiation of sex differentiation in insects is regulated by primary sex determination signals.In the Medfly Ceratitis capitata and other tephritids,Maleness-on-the-Y(MoY) is the master gene for male sex determination.However,the primary signal in Zeugodacus cucurbitae(Coquillett),a very destructive tephritid pest across the world,remains ambiguous.In this study,we have isolated and characterized the Medfly MoY homolog in Z.cucurbitae,ZcMoY.ZcMOY protein shows high sequence conservation to its homologs in Bactrocera species.ZcMoY transcription begins and peaks at very early embryonic stages and then becomes undetectable except the testes and heads of day 1 male adults.Silencing ZcMoY in early embryos by RNAi causes abnormal external genitalia and interior reproductive organs,giving rise to intersexes and feminization of XY individuals.The expression pattern and knockdown phenotypes of ZcMoY indicate that ZcMoY plays a key role in regulating sex determination of Z.cucurbitae males.Our findings will help the understanding of sex determination in Z.cucurbitae and facilitate the development of genetic sexing strains in its biological control.

关键词： MoY Zeugodacus cucurbitae sex determination RNA interference reproductive system

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sex determination and maintenance： the role DMRT1 and FOXL2

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Asian Journal of Andrology 2017年第6期19卷 619-624页

作者： Shengsong Huang Leping Ye Haolin Chen Department of Urology Tongji Hospital Tongji University School of Medicine Shanghai 200065 China Department of Pediatric The Second Affiliated Hospital and Yuying Children's Hospital of Wenzhou Medical University Wenzhou 325027 China.

In many species, including mammals, sex determination is genetically based. The sex chromosomes that individuals carry determine sex identity. Although the genetic base of phenotypic sex is determined at the moment of fertilization, the development of testes or ovaries in the bipotential early gonads takes place during embryogenesis. During development, sex determination depends upon very few critical genes. When one of these key genes functions inappropriately, sex reversal may happen. Consequently, an individual＇s sex phenotype may not necessarily be consistent with the sex chromosomes that are present. For some time, it has been assumed that once the fetal choice is made between male and female in mammals, the gonadal sex identity of an individual remains stable. However, recent studies in mice have provided evidence that it is possible for the gonadal sex phenotype to be switched even in adulthood. These studies have shown that two key genes, doublesex and mad-3 related transcription factor 1 （Dmrtl） and forkhead box L2 （Foxl2）, function in a Yin and Yang relationship to maintain the fates of testes or ovaries in adult mammals, and that mutations in either gene might have a dramatic effect on gonadal phenotype. Thus, adult gonad maintenance in addition to fetal sex determination may both be important for the fertility.

关键词： DMRT1 FOXL2 sex determination sex maintenance

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sex determination and dif ferentiation in Aurelia sp.1: the absence of temperature dependence

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Journal of Oceanology and Limnology 2018年第2期36卷 457-464页

作者：刘春胜顾志峰邢孟欣孙云陈四清陈昭廷 State Key Laboratory of Marine Resource Utilization in South China Sea Hainan University Key Laboratory of Tropical Hydrobiogical Technology Ocean College Hainan University Yellow Sea Fisheries Research Institute Chinese Academy of Fishery Sciences

Cnidarians, being regarded as ‘basal' metazoan animals, are considered to have relatively high plasticity in terms of sex reversal. In this study we used an experimental approach to demonstrate sexual diff erentiation and plasticity in benthic polyps and pelagic medusae of A urelia sp.1 maintained at dif ferent temperatures. Results indicated that in A urelia sp.1, sex diff erentiation has been determined at the polyp stage and that all medusae originating from a given polyp are, phenotypically, of the same sex. In addition, the sex of polyps budding from the same clone(either male or female) at dif ferent temperatures appears to be the same as that of the parent. The sex of medusae that had originated from a known-sex polyp was observed to remain the same as that of the parent, irrespective of differences in strobilation or rearing temperatures. These results indicate that the mechanism of sex determination of Aurelia sp.1. is not influenced by prevailing temperature regimes. A comparison of variability in terms of sexual plasticity of A urelia sp.1 with that of Hydrozoa and Anthozoa suggests that species characterized by a free-swimming medusa life stage have a high dispersal potential, which probably results in a lower rate of sex reversal.

关键词： A urelia sp.1 sex determination temperature sex differentiation

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