家蚕幼虫BmToll9基因对肽聚糖和金黄色葡萄球菌的响应表达
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摘要
【目的】Toll信号通路是昆虫中重要的免疫信号通路,其中Toll受体在保持Toll通路的正常免疫应答、抵抗外源病原体中起到关键的作用。本研究旨在探究肽聚糖和金黄色葡萄球菌Staphylococcus aureus对家蚕Bombyx mori Toll受体基因BmToll9-1和BmToll9-2表达的影响。【方法】将革兰氏阳性菌细胞壁主要成分肽聚糖和革兰氏阳性菌金黄色葡萄球菌分别注射感染家蚕5龄第1天幼虫,诱导其发生免疫反应;采用实时荧光定量PCR分析注射后不同感染时间点BmToll9-2和BmToll9-1基因在家蚕幼虫中肠、表皮、脂肪体和丝腺中的相对表达水平。【结果】往家蚕5龄幼虫中注射肽聚糖或金黄色葡萄球菌后,BmToll9-2基因出现了时间和组织的差异性表达。注射肽聚糖和金黄色葡萄球菌均能诱导5龄幼虫中肠BmToll9-2基因的表达上调,注射肽聚糖和金黄色葡萄球菌分别在3和6 h时对基因表达的诱导效果最好,且注射金黄色葡萄球菌比注射肽聚糖对基因表达的诱导效果更好。注射金黄色葡萄球菌能引起5龄幼虫表皮、脂肪体和丝腺中BmToll9-2基因的表达上调,分别于注射后24, 6和24 h时诱导效果最好。注射金黄色葡萄球菌亦能诱导同源的BmToll9-1基因的上调表达。【结论】家蚕幼虫BmToll9基因在肽聚糖或金黄色葡萄球菌注射处理后均能在不同组织中发生上调表达,推测BmToll9基因参与了家蚕对肽聚糖和金黄色葡萄球菌的免疫反应。
【Aim】 Toll receptors of Toll signaling pathway, an important immune signaling pathway of insects, are essential to maintain immune response against invading exogenous microorganisms in insects. This study aims to explore the influence of peptidoglycan(PGN) and Staphylococcus aureus on the expression of the Toll receptor genes BmToll9-1 and BmToll9-2 in the silkworm, Bombyx mori. 【Methods】 PGN, the main component in cell wall of Gram-positive bacteria, and a Gram-positive bacterium S. aureus were respectively injected into the day-1 5 th instar larvae of B. mori to induce the immune response. The relative expression levels of BmToll9-2 and BmToll9-1 in the midgut, epidermis, fat body and silk glands of the larvae at different time points post injection were detected by real-time quantitative PCR. 【Results】 After injection of PGN or S. aureus into the 5 th instar larvae of B. mori, BmToll9-2 was differentially expressed at different time points and in different tissues. Injection of PGN or S. aureus induced up-regulation of the expression of BmToll9-2 in the midgut of the 5 th instar larvae, achieving the best induction effect at 3 and 6 h post injection, respectively, and the induction effect of S. aureus was better than that of PGN. Injection of S. aureus induced up-regulation of the expression of BmToll9-2 in the epidermis, fat body and silk glands of the 5 th instar larvae, achieving the best induction effects at 24, 6, and 24 h post injection, respectively. Injection of S. aureus also induced up-regulation of the expression of the homogenous BmToll9-1. 【Conclusion】 The expression of BmToll9 genes in different tissues of B. mori larvae is up-regulated after injection of PGN or S. aureus, suggesting that BmToll9 genes are involved in the immune response of B. mori against PGN and S. aureus.
【Aim】 Toll receptors of Toll signaling pathway, an important immune signaling pathway of insects, are essential to maintain immune response against invading exogenous microorganisms in insects. This study aims to explore the influence of peptidoglycan(PGN) and Staphylococcus aureus on the expression of the Toll receptor genes BmToll9-1 and BmToll9-2 in the silkworm, Bombyx mori. 【Methods】 PGN, the main component in cell wall of Gram-positive bacteria, and a Gram-positive bacterium S. aureus were respectively injected into the day-1 5 th instar larvae of B. mori to induce the immune response. The relative expression levels of BmToll9-2 and BmToll9-1 in the midgut, epidermis, fat body and silk glands of the larvae at different time points post injection were detected by real-time quantitative PCR. 【Results】 After injection of PGN or S. aureus into the 5 th instar larvae of B. mori, BmToll9-2 was differentially expressed at different time points and in different tissues. Injection of PGN or S. aureus induced up-regulation of the expression of BmToll9-2 in the midgut of the 5 th instar larvae, achieving the best induction effect at 3 and 6 h post injection, respectively, and the induction effect of S. aureus was better than that of PGN. Injection of S. aureus induced up-regulation of the expression of BmToll9-2 in the epidermis, fat body and silk glands of the 5 th instar larvae, achieving the best induction effects at 24, 6, and 24 h post injection, respectively. Injection of S. aureus also induced up-regulation of the expression of the homogenous BmToll9-1. 【Conclusion】 The expression of BmToll9 genes in different tissues of B. mori larvae is up-regulated after injection of PGN or S. aureus, suggesting that BmToll9 genes are involved in the immune response of B. mori against PGN and S. aureus.
引文
Chen KK,2016.Transcriptome Landscape of Midgut Immunity and Functional Study of Peptidoglycan Recognition Protein-S5 in Bombyx mori.PhD Dissertation,Northwest A&F University,Yangling,Shaanxi.[陈康康,2016.家蚕中肠免疫转录组分析及家蚕肽聚糖识别蛋白-S5的功能研究.陕西杨凌:西北农林科技大学博士学位论文]
Cheng TC,Zhang YL,Liu C,Xu PZ,Gao ZH,Xia QY,Xiang ZH,2008.Identification and analysis of Toll-related genes in the domesticated silkworm,Bombyx mori.Dev.Comp.Immunol.,32(5):464-475.
Chintapalli RT,Hillyer JF,2016.Hemolymph circulation in insect flight appendages:physiology of the wing heart and circulatory flow in the wings of the mosquito Anopheles gambiae.J.Exp.Biol.,219:3945-3951.
Cirimotich CM,Dong YM,Garver LS,Sim S,Dimopoulos G,2010.Mosquito immune defenses against Plasmodium infection.Dev.Comp.Immunol.,34(4):387-395.
Eldon E,Kooyer S,D′Evelyn D,Duman M,Lawinger P,Botas J,Bellen H,1994.The Drosophila 18 wheeler is required for morphogenesis and has striking similarities to Toll.Development,120(4):885-899.
Evans JD,Aronstein K,Chen YP,Hetru C,Imler JL,Jiang H,Kanost M,Thompson GJ,Zou Z,Hultmark D,2006.Immune pathways and defence mechanisms in honey bees Apis mellifera.Insect Mol.Biol.,15(5):645-656.
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Iatsenko I,Kondo S,Mengin-Lecreulx D,Lemaitre B,2016.PGRP-SD,an extracellular pattern-recognition receptor,enhances peptidoglycan-mediated activation of the Drosophila Imd pathway.Cell,45(5):1013-1023.
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Li T,Jiang GB,Lv Y,Li L,Zhang L,Fan XL,Chen P,2015.In vitro induction of Bombyx mori haemolymph melanization by three types of microbes and their cell wall ingredients.Acta Sericol.Sin.,41(2):278-285.[李田,蒋贵兵,吕银,李黎,张亮,范晓林,陈萍,2015.3种微生物及其胞壁物质体外诱导家蚕血液黑化反应的观察.蚕业科学,41(2):278-285]
Liao WL,Bu XL,Jiang WY,Liu JS,2017.Influence of lipopolysaccharide on the expression of the Toll-like receptor gene BmToll9-2 in larval Bombyx mori.Acta Entomol.Sin.,60(4):372-378.[廖文丽,卜晓玲,江婉仪,刘吉升,2017.脂多糖对家蚕幼虫Toll样受体基因BmToll9-2表达的影响.昆虫学报,60(4):372-378]
Lin JH,Xia XF,Yu XQ,Shen JH,Li Y,Lin HL,Tang SS,Vasseur L,You MS,2018.Gene expression profiling provides insights into the immune mechanism of Plutella xylostella midgut to microbial infection.Gene,647:21-30.
Liu JS,Kolliopoulou A,Smagghe G,Swevers L,2014.Modulation of the transcriptional response of innate immune and RNAi genes upon exposure to dsRNA and LPS in silkmoth-derived Bm5 cells overexpressing BmToll9-1 receptor.J.Insect Physiol.,66:10-19.
Liu JS,Smagghe G,Swevers L,2013.Transcriptional response of BmToll9-1 and RNAi machinery genes to exogenous dsRNA in the midgut of Bombyx mori.J.Insect Physiol.,59(6):646-654.
Livak KJ,Schmittgen TD,2001.Analysis of relative gene expression data using real-time quantitative PCR and the 2 (-Delta Delta C(T)) method.Methods,25(4):402-408.
Luna C,Wang X,Huang Y,Zhang J,Zheng L,2002.Characterization of four Toll related genes during development and immune responses in Anopheles gambiae.Insect Biochem.Molec.Biol.,32(9):1171-1179.
Mellroth P,Karlsson J,Steiner H,2003.A scavenger function for Drosophila peptidoglycan recognition protein.J.Biol.Chem.,278(9):7059-7064.
Merkling SH,van Rij RP,2013.Beyond RNAi:antiviral defense strategies in Drosophila and mosquito.J.Insect Physiol.,59(2):159-170.
Nonaka S,Kawamura K,Hori A,Salim E,Fukushima K,Nakanishi Y,Kuraishi T,2018.Characterization of Spz5 as a novel ligand for Drosophila Toll-1 receptor.Biochem.Biophys.Res.Commun.,506(3):510-515.
Nüsslein-Volhard C,Lohs-Schardin M,Sander K,Cremer C,1980.A dorso-ventral shift of embryonic primordia in a new maternal-effect mutant of Drosophila.Nature,283(5746):474-476.
Ramirez JL,Dimopoulos G,2010.The Toll immune signaling pathway control conserved anti-dengue defenses across diverse Ae.aegypti strains and against multiple dengue virus serotypes.Dev.Comp.Immunol.,34(6):625-629.
Takeda K,Akira S,2004.TLR signaling pathways.Semin.Immunol.,16(1):3-9.
Tanaka H,Ishibashi J,Fujita K,Nakajima Y,Sagisaka A,Tomimoto K,Suzuki N,Yoshiyama M,Kaneko Y,Iwasaki T,Sunagawa T,Yamaji K,Asaoka A,Mita K,Yamakawa M,2008.A genome-wide analysis of genes and gene families involved in innate immunity of Bombyx mori.Insect Biochem.Molec.Biol.,38(12):1087-1110.
Valanne S,Wang JH,R?met M,2011.The Drosophila Toll signaling pathway.J.Immunol.,186(2):649-656.
Wang CX,2014.Immune pattern of PGRP and Cecropin genes in Bombyx mori midgut and fat body.J.Shanxi Agric.Univ.(Nat.Sci.Ed.),34(2):109-113.[王传旭,2014.PGRP和Cecropin基因在家蚕中肠和脂肪体中的免疫模式研究.山西农业大学学报(自然科学版),34(2):109-113]
Wang GH,Xia QY,Cheng DJ,Duan J,Zhao P,Chen J,Li Z,2008.Reference genes identified in the silkworm Bombyx mori during metamorphism based on oligonucleotide microarray and confirmed by qRT-PCR.Insect Sci.,15(5):405-413.
Wang YW,Lv ZQ,2016.Expression of 6 antimicrobial peptide genes in silkworm,Bombyx mori after oral bacterial infection.J.Northwest A&F Univ.(Nat.Sci.Ed.),44(1):185-191.[王延稳,吕志强,2016.家蚕肠道感染细菌后6中抗菌肽基因表达的变化.西北农林科技大学学报(自然科学版),44(1):185-191]
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Wu S,Zhang X,He Y,Shuai J,Chen X,Ling E,2010b.Expression of antimicrobial peptide genes in Bombyx mori gut modulated by oral bacterial infection and development.Dev.Comp.Immunol.,34(11):1191-1198.
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Zhang RN,Ren FF,Zhou CB,Xu JF,Yi HY,Ye MQ,Deng XJ,Cao Y,Yu XQ,Yang W,2018.An ML protein from the silkworm Bombyx mori may function as a key accessory protein for lipopolysaccharide signaling.Dev.Comp.Immunol.,88:94-103.
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Cheng TC,Zhang YL,Liu C,Xu PZ,Gao ZH,Xia QY,Xiang ZH,2008.Identification and analysis of Toll-related genes in the domesticated silkworm,Bombyx mori.Dev.Comp.Immunol.,32(5):464-475.
Chintapalli RT,Hillyer JF,2016.Hemolymph circulation in insect flight appendages:physiology of the wing heart and circulatory flow in the wings of the mosquito Anopheles gambiae.J.Exp.Biol.,219:3945-3951.
Cirimotich CM,Dong YM,Garver LS,Sim S,Dimopoulos G,2010.Mosquito immune defenses against Plasmodium infection.Dev.Comp.Immunol.,34(4):387-395.
Eldon E,Kooyer S,D′Evelyn D,Duman M,Lawinger P,Botas J,Bellen H,1994.The Drosophila 18 wheeler is required for morphogenesis and has striking similarities to Toll.Development,120(4):885-899.
Evans JD,Aronstein K,Chen YP,Hetru C,Imler JL,Jiang H,Kanost M,Thompson GJ,Zou Z,Hultmark D,2006.Immune pathways and defence mechanisms in honey bees Apis mellifera.Insect Mol.Biol.,15(5):645-656.
Gerardo NM,Altincicek B,Anselme C,Atamian H,Barribeau SM,de Vos M,Duncan EJ,Evans JD,Gabaldón T,Ghanim M,Heddi A,Kaloshian I,Latorre A,Moya A,Nakabachi A,Parker BJ,Pérez-Brocal V,Pignatelli M,Rahbé Y,Ramsey JS,Spragg CJ,Tamames J,Tamarit D,Tamborindeguy C,Vincent-Monegat C,Vilcinskas A,2011.Immunity and other defenses in pea aphids,Acyrthosiphon pisum.Genome Biol.,11(2):R21.
Hetru C,Hoffmann JA,2009.NF-kappaB in the immune response of Drosophila.Cold Spring Harb.Perspect.Biol.,1(6):a000232.
Iatsenko I,Kondo S,Mengin-Lecreulx D,Lemaitre B,2016.PGRP-SD,an extracellular pattern-recognition receptor,enhances peptidoglycan-mediated activation of the Drosophila Imd pathway.Cell,45(5):1013-1023.
Kanost MR,Arrese EL,Cao XL,Chen YR,Chellapilla S,Goldsmith MR,Grosse-Whilde E,Heckel DG,Herndon N,Jiang HB,Papanicolaou A,Qu JX,Soulages JL,Vogel H,Walters J,Waters J,Waterhouse RM,Ahn SJ,Almeida FC,An CJ,Aqrawi P,Bretschneider A,Bryant WB,Bucks S,Chao H,Chevignon G,Christen JM,Clarke DF,Dittmer NT,Ferguson LCF,Garacelous S,Gordon KHJ,Gunaratna RT,Han Y,Hauser F,He Y,Heidel-Fischer H,Hirsh A,Hu YX,Jiang HB,Kalra D,Klinner C,K?nig C,Kovar C,Kroll AR,Kuwar SS,Lee SL,Lehman R,Li K,Li ZF,Liang HQ,lovelace S,Lu ZQ,Mansfield JH,McCulloch KJ,Mathew T,Morton B,Muzny DM,Neunemann D,Ongeri F,Pauchet Y,Pu LL,Pyrousis I,Rao XJ,Redding A,Roesel C,Sanchez-Gracia A,Schaack S,Shuakla A,Tetreau G,Wang Y,Xiong GH,Traut W,Walsh TK,Worley KC,Wu D,Wu WB,Wu YQ,Zhang XF,Zou Z,Zucker H,Briscoe AD,Burmester T,Clem RJ,Feyereisen R,Grimmelikhuijizen CJP,Hamodrakas SJ,Hansson BS,Huguet E,Jermiin LS,Lan Q,Lehman HK,Lorenzen M,Merzendorfer H,Michalopoulos I,Morton DB,Muthukrishman S,Oakeshott JG,Palmer W,Park Y,Passarelli AL,Rozas J,Schwartz LM,Smith W,Southgate A,Vilcinskas A,Vogt R,Wang P,Werren J,Yu XQ,Zhou JJ,Brown SJ,Scherer SE,Richards S,Blissard GW,2016.Multifaceted biological insights from a draft genome sequence of the tobacco hornworm moth,Manduca sexta.Insect Biochem Molec.Biol.,76:118-147.
Li T,Jiang GB,Lv Y,Li L,Zhang L,Fan XL,Chen P,2015.In vitro induction of Bombyx mori haemolymph melanization by three types of microbes and their cell wall ingredients.Acta Sericol.Sin.,41(2):278-285.[李田,蒋贵兵,吕银,李黎,张亮,范晓林,陈萍,2015.3种微生物及其胞壁物质体外诱导家蚕血液黑化反应的观察.蚕业科学,41(2):278-285]
Liao WL,Bu XL,Jiang WY,Liu JS,2017.Influence of lipopolysaccharide on the expression of the Toll-like receptor gene BmToll9-2 in larval Bombyx mori.Acta Entomol.Sin.,60(4):372-378.[廖文丽,卜晓玲,江婉仪,刘吉升,2017.脂多糖对家蚕幼虫Toll样受体基因BmToll9-2表达的影响.昆虫学报,60(4):372-378]
Lin JH,Xia XF,Yu XQ,Shen JH,Li Y,Lin HL,Tang SS,Vasseur L,You MS,2018.Gene expression profiling provides insights into the immune mechanism of Plutella xylostella midgut to microbial infection.Gene,647:21-30.
Liu JS,Kolliopoulou A,Smagghe G,Swevers L,2014.Modulation of the transcriptional response of innate immune and RNAi genes upon exposure to dsRNA and LPS in silkmoth-derived Bm5 cells overexpressing BmToll9-1 receptor.J.Insect Physiol.,66:10-19.
Liu JS,Smagghe G,Swevers L,2013.Transcriptional response of BmToll9-1 and RNAi machinery genes to exogenous dsRNA in the midgut of Bombyx mori.J.Insect Physiol.,59(6):646-654.
Livak KJ,Schmittgen TD,2001.Analysis of relative gene expression data using real-time quantitative PCR and the 2 (-Delta Delta C(T)) method.Methods,25(4):402-408.
Luna C,Wang X,Huang Y,Zhang J,Zheng L,2002.Characterization of four Toll related genes during development and immune responses in Anopheles gambiae.Insect Biochem.Molec.Biol.,32(9):1171-1179.
Mellroth P,Karlsson J,Steiner H,2003.A scavenger function for Drosophila peptidoglycan recognition protein.J.Biol.Chem.,278(9):7059-7064.
Merkling SH,van Rij RP,2013.Beyond RNAi:antiviral defense strategies in Drosophila and mosquito.J.Insect Physiol.,59(2):159-170.
Nonaka S,Kawamura K,Hori A,Salim E,Fukushima K,Nakanishi Y,Kuraishi T,2018.Characterization of Spz5 as a novel ligand for Drosophila Toll-1 receptor.Biochem.Biophys.Res.Commun.,506(3):510-515.
Nüsslein-Volhard C,Lohs-Schardin M,Sander K,Cremer C,1980.A dorso-ventral shift of embryonic primordia in a new maternal-effect mutant of Drosophila.Nature,283(5746):474-476.
Ramirez JL,Dimopoulos G,2010.The Toll immune signaling pathway control conserved anti-dengue defenses across diverse Ae.aegypti strains and against multiple dengue virus serotypes.Dev.Comp.Immunol.,34(6):625-629.
Takeda K,Akira S,2004.TLR signaling pathways.Semin.Immunol.,16(1):3-9.
Tanaka H,Ishibashi J,Fujita K,Nakajima Y,Sagisaka A,Tomimoto K,Suzuki N,Yoshiyama M,Kaneko Y,Iwasaki T,Sunagawa T,Yamaji K,Asaoka A,Mita K,Yamakawa M,2008.A genome-wide analysis of genes and gene families involved in innate immunity of Bombyx mori.Insect Biochem.Molec.Biol.,38(12):1087-1110.
Valanne S,Wang JH,R?met M,2011.The Drosophila Toll signaling pathway.J.Immunol.,186(2):649-656.
Wang CX,2014.Immune pattern of PGRP and Cecropin genes in Bombyx mori midgut and fat body.J.Shanxi Agric.Univ.(Nat.Sci.Ed.),34(2):109-113.[王传旭,2014.PGRP和Cecropin基因在家蚕中肠和脂肪体中的免疫模式研究.山西农业大学学报(自然科学版),34(2):109-113]
Wang GH,Xia QY,Cheng DJ,Duan J,Zhao P,Chen J,Li Z,2008.Reference genes identified in the silkworm Bombyx mori during metamorphism based on oligonucleotide microarray and confirmed by qRT-PCR.Insect Sci.,15(5):405-413.
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