斑翅果蝇气味结合蛋白OBP56h与小分子化合物的结合特征
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摘要
【目的】克隆斑翅果蝇(Drosophila suzukii)气味结合蛋白56h(odorant binding protein 56h,OBP56h)基因,诱导表达斑翅果蝇OBP56h重组蛋白(DsuzOBP56h),研究其与小分子化合物的结合特性。【方法】通过RT-PCR并设计特异性引物克隆斑翅果蝇OBP56h ORF全长,从NCBI数据库中下载相似度高的昆虫气味结合蛋白序列,进行序列比对和分析。以NdeⅠ和XhoⅠ为酶切位点,将OBP56h连入pET-30a(+)原核表达载体,将重组质粒转入BL21(DE3)大肠杆菌感受态细胞。扩大培养阳性菌株,并用IPTG诱导表达DsuzOBP56h重组蛋白。收集菌液,通过超声波破碎细胞得到蛋白,利用Ni-NTA柱纯化蛋白,进行Tris-HCl透析,用BCA法测定蛋白浓度。蛋白用50 mmol·L~(-1) Tris-HCl(pH 7.4)稀释至终浓度2μmol·L~(-1),配基用色谱级甲醇稀释至终浓度1 mmol·L~(-1),以4,4′-二苯胺基-1,1′-联萘-5,5′-二磺酸二钾盐(4,4′-dianilino-1,1′-binaphthyl-5,5′-disulfonicacid dipotassium salt,bis-ANS)荧光探针为报告子,利用荧光竞争结合试验检测DsuzOBP56h蛋白与18种候选小分子化合物配基的结合特性。【结果】克隆获得了斑翅果蝇OBP56h的ORF全长,共405 bp,N-端含有19个氨基酸组成的信号肽,具有6个保守半胱氨酸位点,符合OBP的典型特征,与其同属的黑腹果蝇OBP56h进化关系最近。成功连入pET-30a(+)表达载体,在1 mmol·L~(-1) IPTG、28℃条件下诱导DsuzOBP56h蛋白表达,并过柱纯化得到目的蛋白。荧光光谱试验显示,荧光探针bis-ANS与DsuzOBP56h的解离常数为0.9568μmol·L~(-1),适合作为本试验中竞争性荧光结合试验的报告子;进一步的荧光竞争结合试验表明,在18种候选配基中,苦味物质盐酸小蘖碱和香豆素与DsuzOBP56h的结合亲和性较强,解离常数分别为12.16和17.93μmol·L~(-1),柚皮素与DsuzOBP56h的解离常数为25.32μmol·L~(-1),草莓叶片产生的一种对斑翅果蝇具有吸引作用的挥发性气味物质β-环柠檬醛也能与DsuzOBP56h结合,其解离常数为31.37μmol·L~(-1)。【结论】斑翅果蝇气味结合蛋白OBP56h能与测试的多种植物苦味物质和挥发性气味物质结合,表明DsuzOBP56h很有可能参与斑翅果蝇对食物味觉和嗅觉的识别行为,研究结果可为理解斑翅果蝇的取食行为提供理论依据,并为开展斑翅果蝇的生态防控提供新思路。
【Objective】The objective of this study is to clone the odorant binding protein 56 h(OBP56 h) gene from Drosophila suzukii, get the recombinant DsuzOBP56 h protein, and characterize the binding profiles of DsuzOBP56 h with some small molecular compounds.【Method】By means of specific primer, reverse transcription PCR(RT-PCR) was used to clone the full-length ORF of Dsuz OBP56 h. The sequences of insect OBPs with high similarity were downloaded from NCBI database for sequence alignment and analysis. Using NdeⅠand XhoⅠas restriction sites, OBP56 h was ligated into pET-30 a(+) prokaryotic expression vector, and transformed into Escherichia coli BL21(DE3). IPTG was applied to induce the expression of recombinant Dsuz OBP56 h protein.The bacterial solution was collected and the protein was obtained through breaking cells by ultrasound, and then the protein was purified by the method of Ni-NTA resin. The purified protein was dialyzed by Tris-HCl and the concentration was determined by the method of BCA. The protein was diluted with 50 mmol·L-1 Tris-HCl(pH 7.4) to a final concentration of 2 μmol·L-1, and the ligand was diluted with chromatography-grade methanol to a final concentration of 1 mmol·L-1. The binding characterization of DsuzOBP56 h with 18 small molecular compounds was investigated using bis-ANS as fluorescence probe. 【Result】 The full-length ORF of OBP56 h in D. suzukii was amplified, which is 405 bp in total, including 19 amino acids of signal peptide in the N-terminal. It has 6 conserved cysteine sites, which is consistent with the typical characteristics of OBPs, and has the closest evolutionary relationship with D. melanogaster OBP56 h. OBP56 h was successfully inserted into pET-30 a(+) and expressed at 1 mmol·L-1 IPTG and 28℃,then purified by the method of Ni-NTA resin. In the competitive fluorescence assay, the dissociation constant Kbis-ANS was 0.9568μmol·L-1, indicating that bis-ANS is suitable to be a reporter of competitive fluorescence binding assay. Among 18 ligands, the binding affinity of bitter tastants berberine chloride and coumarin with DsuzOBP56 h was strong, and the dissociation constant is12.16 and 17.93 μmol·L-1, respectively. The dissociation constant of naringenin with DsuzOBP56 h is 25.32 μmol·L-1. A volatile odorant β-cyclocitral, which is attractive to D. suzukii produced by strawberry leaves, can also bind to DsuzOBP56 h, with the dissociation constant of 31.37 μmol·L-1.【Conclusion】The OBP56 h in D. suzukii can bind with a variety of bitter tastants and volatile odors from plants, indicating that OBP56 h may be involved in the gustatory and olfactory recognition of food in D. suzukii.The results can provide a theoretical basis for understanding the feeding behavior of D. suzukii, and provide a new idea for the ecological prevention and control of D. suzukii.
【Objective】The objective of this study is to clone the odorant binding protein 56 h(OBP56 h) gene from Drosophila suzukii, get the recombinant DsuzOBP56 h protein, and characterize the binding profiles of DsuzOBP56 h with some small molecular compounds.【Method】By means of specific primer, reverse transcription PCR(RT-PCR) was used to clone the full-length ORF of Dsuz OBP56 h. The sequences of insect OBPs with high similarity were downloaded from NCBI database for sequence alignment and analysis. Using NdeⅠand XhoⅠas restriction sites, OBP56 h was ligated into pET-30 a(+) prokaryotic expression vector, and transformed into Escherichia coli BL21(DE3). IPTG was applied to induce the expression of recombinant Dsuz OBP56 h protein.The bacterial solution was collected and the protein was obtained through breaking cells by ultrasound, and then the protein was purified by the method of Ni-NTA resin. The purified protein was dialyzed by Tris-HCl and the concentration was determined by the method of BCA. The protein was diluted with 50 mmol·L-1 Tris-HCl(pH 7.4) to a final concentration of 2 μmol·L-1, and the ligand was diluted with chromatography-grade methanol to a final concentration of 1 mmol·L-1. The binding characterization of DsuzOBP56 h with 18 small molecular compounds was investigated using bis-ANS as fluorescence probe. 【Result】 The full-length ORF of OBP56 h in D. suzukii was amplified, which is 405 bp in total, including 19 amino acids of signal peptide in the N-terminal. It has 6 conserved cysteine sites, which is consistent with the typical characteristics of OBPs, and has the closest evolutionary relationship with D. melanogaster OBP56 h. OBP56 h was successfully inserted into pET-30 a(+) and expressed at 1 mmol·L-1 IPTG and 28℃,then purified by the method of Ni-NTA resin. In the competitive fluorescence assay, the dissociation constant Kbis-ANS was 0.9568μmol·L-1, indicating that bis-ANS is suitable to be a reporter of competitive fluorescence binding assay. Among 18 ligands, the binding affinity of bitter tastants berberine chloride and coumarin with DsuzOBP56 h was strong, and the dissociation constant is12.16 and 17.93 μmol·L-1, respectively. The dissociation constant of naringenin with DsuzOBP56 h is 25.32 μmol·L-1. A volatile odorant β-cyclocitral, which is attractive to D. suzukii produced by strawberry leaves, can also bind to DsuzOBP56 h, with the dissociation constant of 31.37 μmol·L-1.【Conclusion】The OBP56 h in D. suzukii can bind with a variety of bitter tastants and volatile odors from plants, indicating that OBP56 h may be involved in the gustatory and olfactory recognition of food in D. suzukii.The results can provide a theoretical basis for understanding the feeding behavior of D. suzukii, and provide a new idea for the ecological prevention and control of D. suzukii.
引文
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[2]ROTA-STABELLI O,BLAXTER M,ANFORA G.Drosophila suzukii.Current Biology,2013,23(1):R8-R9.
[3]LEAL W S.Odorant reception in insects:Roles of receptors,binding proteins,and degrading enzymes.Annual Review of Entomology,2013,58:373-391.
[4]SWARUP S,WILLIAMS T I,ANHOLT R R.Functional dissection of odorant binding protein genes in Drosophila melanogaster.Genes,Brain,and Behavior,2011,10(6):648-657.
[5]WANG P,LYMAN R F,MACKAY T F,ANHOLT R R.Natural variation in odorant recognition among odorant-binding proteins in Drosophila melanogaster.Genetics,2010,184(3):759-767.
[6]GRO?E-WILDE E,SVATO?A,KRIEGER J.A pheromone-binding protein mediates the bombykol-induced activation of a pheromone receptor in vitro.Chemical Senses,2006,31(6):547-555.
[7]陈玲,李红亮,周宇翔,赵磊,张林雅,倪翠侠,商晗武.桔小实蝇气味结合蛋白BdorOBP2的c DNA克隆、组织表达及配基结合特性.昆虫学报,2013,56(6):612-621.CHEN L,LI H L,ZHOU Y X,ZHAO L,ZHANG L Y,NI C X,SHANG H W.c DNA cloning,tissue expression and ligand binding characteristics of odorant-binding protein 2 from the oriental fruit fly,Bactrocera dorsalis(Diptera:Tephritidae).Acta Entomologica Sinica,2013,56(6):612-621.(in Chinese)
[8]陈东凯,张林雅,邢振龙,雷仲仁.美洲斑潜蝇气味结合蛋白OBP13的鉴定与功能.中国农业科学,2018,51(5):893-904.CHEN D K,ZHANG L Y,XING Z L,LEI Z R.Identification and function of the OBP13 protein from the leafminer(Liriomyza sativae).Scientia Agricultura Sinica,2018,51(5):893-904.(in Chinese)
[9]KARAGEORGI M,BR?CKER L B,LEBRETON S,MINERVINOC,CAVEY M,SIJU K P,GRUNWALD KADOW I C,GOMPEL N,PRUD’HOMME B.Evolution of multiple sensory systems drives novel egg-laying behavior in the fruit pest Drosophila suzukii.Current Biology,2017,27(6):847-853.
[10]任路明,秦胜楠,丁心婷,万方浩,褚栋.水果害虫铃木氏果蝇的入侵及其防控研究进展.生物安全学报,2014,23(3):142-150.REN L M,QIN S N,DING X T,WAN F H,CHU D.Research on the invasion and control of a fruit insect pest,Drosophila suzukii(Matsumura).Journal of Biosafety,2014,23(3):142-150.(in Chinese)
[11]张开春,闫国华,郭晓军,王晶,张晓明,周宇.斑翅果蝇(Drosophila suzukii)研究现状.果树学报,2014,31(4):717-721.ZHANG K C,YAN G H,GUO X J,WANG J,ZHANG X M,ZHOUY.Research review on spotted wing drosophila(Drosophila suzukii).Journal of Fruit Science,2014,31(4):717-721.(in Chinese)
[12]HAMBY K A,BECHER P G.Current knowledge of interactions between Drosophila suzukii and microbes,and their potential utility for pest management.Journal of Pest Science,2016,89(3):621-630.
[13]LEE J C,BURRACK H J,BARRANTES L D,BEERS E H,DREVES A J,HAMBY K A,HAVILAND D R,ISAACS R,RICHARDSON T A,SHEARER P W,STANLEY C A,WALSH D B,WALTON V M,ZALOM F G,BRUCK D J.Evaluation of monitoring traps for Drosophila suzukii(Diptera:Drosophilidae)in North America.Journal of Economic Entomology,2012,105(4):1350-1357.
[14]WANG X G,STEWART T J,BIONDI A,CHAVEZ B A,INGELS C,CAPRILE J,GRANT J A,WALTON V M,DAANE K M.Population dynamics and ecology of Drosophila suzukii in central California.Journal of Pest Science,2016,89(3):701-712.
[15]FENG Y,BRUTON R,PARK A,ZHANG A.Identification of attractive blend for spotted wing drosophila,Drosophila suzukii,from apple juice.Journal of Pest Science,2018,91(4):1251-1267.
[16]WALLINGFORD A K,HESLER S P,CHA D H,LOEB G M.Behavioral response of spotted-wing drosophila,Drosophila suzukii Matsumura,to aversive odors and a potential oviposition deterrent in the field.Pest Management Science,2016,72(4):701-706.
[17]GALINDO K,SMITH D P.A large family of divergent Drosophila odorant-binding proteins expressed in gustatory and olfactory sensilla.Genetics,2001,159(3):1059-1072.
[18]SWARUP S,MOROZOVA T V,SRIDHAR S,NOKES M,ANHOLTR R.Modulation of feeding behavior by odorant-binding proteins in Drosophila melanogaster.Chemical Senses,2014,39(2):125-132.
[19]DALTON D T,WALTON V M,SHEARER P W,WALSH D B,CAPRILE J,ISAACS R.Laboratory survival of Drosophila suzukii under simulated winter conditions of the Pacific Northwest and seasonal field trapping in five primary regions of small and stone fruit production in the United States.Pest Management Science,2011,67(11):1368-1374.
[20]CONGDON R W,MUTH G W,SPLITTGERBER A G.The binding interaction of Coomassie blue with proteins.Analytical Biochemistry,1993,213(2):407-413.
[21]KNOLHOFF L M,HECKEL D G.Behavioral assays for studies of host plant choice and adaptation in herbivorous insects.Annual Review of Entomology,2014,59:263-278.
[22]MATSUO T,SUGAYA S,YASUKAWA J,AIGAKI T,FUYAMA Y.Odorant-binding proteins OBP57d and OBP57e affect taste perception and host-plant preference in Drosophila sechellia.PLoS Biology,2007,5(5):e118.
[23]张帅,张永军,苏宏华,高希武,郭予元.中红侧沟茧蜂化学感受蛋白MmedCSP1的结合特征.昆虫学报,2009,52(8):838-844.ZHANG S,ZHANG Y J,SU H H,GAO X W,GUO Y Y.Binding characterization of chemosensory protein MmedCSP1 in Microplitis mediator(Hymenoptera:Braconidae).Acta Entomologica Sinica,2009,52(8):838-844.(in Chinese)
[24]L?BEL D,STROTMANN J,JACOB M,BREER H.Identification of a third rat odorant-binding protein(OBP3).Chemical Senses,2001,26(6):673-680.
[25]XU P,ATKINSON R,JONES D N,SMITH D P.Drosophila OBPLUSH is required for activity of pheromone-sensitive neurons.Neuron,2005,45(2):193-200.
[26]LAUGHLIN J D,HA T S,JONES D N,SMITH D P.Activation of pheromone-sensitive neurons is mediated by conformational activation of pheromone-binding protein.Cell,2008,133(7):1255-1265.
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