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2586 CUI Dong-nan et al. Journal of Integrative Agriculture 2019, 18(11): 2579–2588 acyl-CoAs. In our study, we observed the significant down- regulation of peroxisomal multifunctional enzyme type 2 isoform ×1 in diapause eggs. Fatty acyl-CoA reductase 1, a critical enzyme in plasmalogen biosynthesis, can reduce fatty acids into their respective fatty alcohols (Cheng and Russell 2004; Honsho et al. 2013). The significant increased expression of five genes encoding the fatty acyl-CoA reductase 1 has been observed in oocytes of Bombyx mori producing diapause eggs (Chen et al. 2017). The putative fatty acyl-CoA reductase was up-regulated in diapause eggs in this study. However, the difference was insignificant. Additionally, superoxide dismutase (Cu-Zn)-like precursor and superoxide dismutase (Mn), two typical antioxidant enzymes were less abundant in diapause eggs than that in non-diapause eggs but with no significant difference. Similar results were also observed in diapause flesh fly, Sarcophaga crassipalpis , and bivoltine silkworm, B . mori (Ragland et al. 2010; Chen et al. 2017). 5. Conclusion Through proteome analysis of both diapause and non- diapause eggs, we found that maternal effects may regulate diapause through hormone biosynthesis, the longevity regulating pathway and peroxisome pathway. Our study could be informative for further future research on the mechanisms of diapause induced by maternal effects in migratory locust. Acknowledgements This research was funded by the earmarked fund of China Agriculture Research System (CARS-34-07), and the National Natural Science Foundation of China (31672485). We thank anonymous reviewers for helpful suggestions on an earlier draft of this manuscript. References Bradford M M. 1976. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Analytical Biochemistry , 72 , 248–254. Chapuis M P, Simpson S J, Blondin L, Sword G A. 2011. Taxa-specific heat shock proteins are over-expressed with crowding in the Australian plague locust. Journal of Insect Physiology , 57 , 1562–1567. Chen Y R, Jiang T, Zhu J, Xie Y C, Tan Z C, Chen Y H, Tang S M, Hao B F, Wang S P, Huang J S, Shen X J. 2017. Transcriptome sequencing reveals potential mechanisms of diapause preparation in bivoltine silkworm Bombyx mori (Lepidoptera: Bombycidae). Comparative Biochemistry and Physiology (Part D: Genomics and Proteomics), 24 , 68–78. Cheng J B, Russell D W. 2004. Mammalian wax biosynthesis I. Identification of two fatty acyl-coenzyme a reductases with different substrate specificities and tissue distributions. Journal of Biological Chemistry , 279 , 37789–37797. Deng Y, Li F, Rieske L K, Sun L L, Sun S H. 2018. Transcriptome sequencing for identification of diapause-associated genes in fall webworm, Hyphantria cunea Drury. Gene , 668 , 229–236. Denlinger D L. 2002. Regulation of diapause. Annual Review of Entomology , 47 , 93–122. Denlinger D L, Armbruster P A. 2014. Mosquito diapause. Annual Review of Entomology , 59 , 73–93. Gilbert L I, Warren J T. 2005. A molecular genetic approach to the biosynthesis of the insect steroid molting hormone. Vitamins and Hormones , 73 , 31–57. Godlewski J, Kludkiewicz B, Grzelak K, Cymborowski B. 2001. Expression of larval hemolymph proteins ( Lhp ) genes and protein synthesis in the fat body of greater wax moth ( Galleria mellonella ) larvae during diapause. Journal of Insect Physiology , 47 , 759–766. Guo F, Chen Y L, Lu B L. 1991. Biology of Chinese Migratory Locust . Shandong Science and Technology Press, China. p. 320. (in Chinese) Guo Y L, Pang S T, Ahmed U A E, Hao Z P, Shi Z H. 2007. Effect of maternal photoperiod experience on diapause incidence of Cotesia plutellae . Chinese Journal of Biological Control , 23 , 1–4. (in Chinese) Han B, Fang Y, Feng M, Hu H, Hao Y, Ma C, Huo X M, Meng L F, Zhang X F, Wu F, Li J K. 2017. Brain membrane proteome and phosphoproteome reveal molecular basis associating with nursing and foraging behaviors of honeybee workers. Journal of Proteome Research , 16 , 3646–3663. Hao K, Wang J, Tu X B, Whitman D W, Zhang Z H. 2017. Transcriptomic and proteomic analysis of Locusta migratoria * * * * 0 0.3 0.6 0.9 1.2 1.5 Hexamerin-like protein 4 Juvenile hormone epoxide hydrolase 1 Cytochrome P450 Heat shock protein 20.7 Relative mRNA level Non-diapause eggs Diapause eggs Fig. 5 The relative quantitative expression of four specific proteins using 2 –ΔΔC T methods. * indicates a significant difference of mRNA expression between non-diapause eggs and diapause eggs (independent sample t -test, P <0.05).