书虱谷胱甘肽S-转移酶的纯化及生化毒理学特性研究
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摘要
书虱隶属于啮目Psocoptera、书虱科Liposcelididae、书虱属Liposcelis,是储藏物中常见的害虫。书虱属昆虫在世界各地广为分布,目前已知种类123种,我国共记载有27种。作为储粮生态中的害虫优势种群,嗜卷书虱Liposcelis bostrychophila Badonnel和嗜虫书虱L.entomophila(Enderlein)经常混合发生,造成严重的经济损失,而且对化学药剂的抗性发展很快,已经引起了全世界储藏物工作者的高度重视。
蛋白质分离和纯化是进行蛋白质结构和功能研究的基础。采用蛋白质纯化的方法分离参与抗药性产生的酶,明确其生物化学特性及其与抗药性的关系,对于害虫抗药性治理具有重要的意义。谷胱甘肽S-转移酶(Glutathione S-transferases,GSTs,EC 2.5.1.18)是一类多功能的超基因家族酶,广泛分布于哺乳动物、昆虫、植物以及微生物等生物体内。这类酶能催化内源还原性谷胱甘肽(GSH)与各种有害的亲电性底物相结合,增加后者的可溶性从而有利于其从细胞内排出,进而保护生物体内的核酸和蛋白质免受亲电基团攻击。同时,它也参与激素的胞内运输、合成,清除外源化合物所产生的有害的氧自由基以及保护细胞免遭氧化压力胁迫。本学位论文在教育部新世纪优秀人才支持计划(NCET-04-0854)和教育部高等学校博士点专项基金(200806350009)的资助下,利用亲和层析法对嗜卷书虱和嗜虫书虱的GSTs进行分离纯化的基础上,对纯化产物的生化毒理学特性进行了较为全面的解析,研究结果为揭示GSTs介导的书虱适应环境胁迫的机理提供了生化毒理学依据,同时丰富和发展了储藏物害虫抗性机理研究的科学理论体系。主要研究结果总结如下:
1嗜卷书虱不同品系GSTs的纯化及生化毒理学特性解析
采用Glutathione Sepharose 4B亲和层析法纯化了嗜卷书虱3个品系(SS,敏感品系;DDVP-R,敌敌畏抗性品系;PH_3-R,磷化氢抗性品系)的GSTs,纯化产物达电泳级纯度。经SDS-PAGE电泳检测,纯化后GSTs的蛋白分子量约为23 kDa。利用Edman降解法对纯化产物N端前15个氨基酸测序获得的短肽为:APKYKYL TYPFNSILTGLGL,该序列与Sigma类昆虫GSTs N端氨基酸序列高度一致,表明纯化获得的GSTs属于昆虫Sigma类。
嗜卷书虱3个品系GSTs的纯化倍数在62-91倍之间,回收率介于27-41%。利用微量滴度酶标板法对纯化后的GSTs的生化特性分析表明,与SS品系相比,DDVP-R和PH_3-R品系的GSTs活性显著提高(P＜0.05),而两抗性品系间相比较差异未达显著水平(P＞0.05)。进一步的动力学比较研究发现,以CDNB为底物时,DDVP-R品系的K_m值最高;而以GSH为底物时,两抗性品系K_m值较低,说明DDVP和PH_3的胁迫选择造成GSTs对内源性化合物GSH获得了较高的亲和能力;对V_(max)而言,两抗性品系的V_(max)值均高于敏感品系,说明其催化能力提高。
嗜卷书虱3个品系GSTs活力随pH值变化的趋势大体相同,其中SS品系GSTs的最适pH范围较广,介于pH7.0-pH8.0之间,而两抗性品系的最适范围介于pH 7.0-pH 7.5之间。酶促反应最适温度研究发现,SS和PH_3-R品系在35℃时GSTs活力最高,而DDVP-R品系则在40℃时GSTs的活力最高。
杀虫剂对嗜卷书虱3个品系GSTs的离体作用研究表明,供试的杀虫剂中丁硫克百威对嗜卷书虱3个品系的GSTs的抑制作用明显,3品系间相比较PH_3-R品系GSTs对丁硫克百威的抑制作用敏感度低,抑制中浓度I_(50)分别为SS和DDVP-R品系的3.9和4.7倍。毒死蜱和高效氯氰菊酯对嗜卷书虱的GSTs仅有部分的抑制作用,毒死蜱在低浓度时还表现为一定的激活作用。利尿酸、姜黄素和四溴磺酚钠对嗜卷书虱纯化后的GSTs均有显著的离体抑制作用。其中,姜黄素的抑制作用最强,其I_(50)达纳摩尔数量级。综合分析,SS品系与两抗性品系相比对上述抑制剂更为敏感,两抗性品系GSTs对各抑制剂的敏感性存在一定的差异。
2嗜卷书虱不同发育阶段GSTs的纯化及生化毒理学特性解析
采用Glutathione Sepharose 4B亲和层析法纯化了嗜卷书虱3个发育阶段(1-2龄低龄若虫、3-4龄高龄若虫和成虫)的GSTs。结果表明,低龄若虫GSTs的纯化倍数最高,达102倍,但回收率仅为28%;高龄若虫GSTs的纯化倍数最低,仅为63倍,但回收率高达61%;成虫GSTs的纯化倍数介于低龄和高龄若虫之间,其GSTs的比活力最高,且显著高于低龄若虫(P＜0.05)。以GSH和CDNB为底物的动力学特性分析发现,低龄若虫GSTs的K_m值最低,而成虫GSTs对两底物均表现最高的催化反应速度V_(max),表明成虫期GSTs对底物的催化能力显著高于若虫期。
毒死蜱对嗜卷书虱不发育阶段GSTs纯化后的离体作用结果表明,该药剂对GSTs有一定的抑制作用,但在低浓度时表现为一定的激活作用,随着浓度的升高,抑制作用逐渐显现。在测试的5个浓度下,毒死蜱对GSTs的最高抑制率仅为39%。5种典型化合物(利尿酸、姜黄素、双硫仑、马来酸二乙酯和四溴磺酚钠)对嗜卷书虱各发育阶段GSTs均有显著的离体抑制作用,其中GSTs对双硫仑的耐受性最强,该抑制剂的离体I_(50)达毫摩尔数量级。综合分析发现,除高龄若虫对利尿酸最不敏感外,嗜卷书虱成虫对其它抑制剂均表现为最不敏感。
此外,研究了金属离子(Zn~(2+)、Hg~(2+)、Mn~(2+)、Cu~(2+)和Ca~(2+))对嗜卷书虱GSTs活性的影响。结果表明,5种金属离子对嗜卷书虱GSTs均有显著的抑制作用。嗜卷书虱各发育阶段的GSTs对Ca~(2+)最不敏感,I_(50)达纳摩尔数量级;Hg~(2+)对嗜卷书虱GSTs的离体抑制作用最为显著,但各虫态间I_(50)值无显著性差异(P＞0.05)。
3嗜卷书虱不同地理种群GSTs的纯化及生化毒理学特性解析
采用Glutathione Sepharose 4B亲和层析法纯化了嗜卷书虱3个地理种群(四川广汉、四川简阳和实验室保存种群)的GSTs。结果表明,经亲和层析后,实验室种群GSTs的纯化倍数高达121倍,回收率亦最高,达52%;四川广汉种群纯化倍数和回收率分别为58和33%;简阳种群的纯化倍数最低,仅为40倍,回收率仅为16%。总蛋白和活力测定结果显示,实验室种群GSTs纯化产物的总蛋白含量和酶活力显著高于其余两个种群(P＜0.05),但其它种群之间各参数均无显著性差异(P＞0.05)。
分别以CDNB和GSH为底物,利用双倒数作图法获得了嗜卷书虱3个种群GSTs纯化后的动力学参数。比较分析K_m值发现,实验室种群对2种底物的亲和力最低,简阳种群对底物GSH的亲和力最高,而广汉种群对底物CDNB的亲和力最高;就V_(max)而言,实验室种群对底物GSH和CDNB均表现出最高的催化活性,且显著高于简阳种群与广汉种群(P＜0.05)。
杀虫剂的离体抑制作用测定结果表明,广汉种群对丁硫克百威最为敏感,简阳种群其次,实验室种群对该药剂的离体抑制作用最不敏感。高效氯氰菊酯对嗜卷书虱不同种群纯化后的GSTs有部分的抑制作用,且各种群受到的离体作用类似。利尿酸、姜黄素、马来酸二乙酯和四溴磺酚钠对嗜卷书虱3个种群均表现为显著的抑制作用,其中广汉种群GSTs对各种抑制剂的作用最为敏感。
各金属离子对嗜卷书虱3个种群的作用结果显示,GSTs对Ca~(2+)最不敏感,其中实验室种群与其它两种群相比更不敏感,其I_(50)值最高;Hg~(2+)对GSTs离体抑制作用最为强烈,各种群间I_(50)值无显著性差异(P＞0.05);Mn~(2+)和Ca~(2+)对各种群GSTs的离体作用较为相似,其中广汉种群对这2种金属离子的抑制作用最为敏感,而实验室种群最不敏感。
4嗜虫书虱不同地理种群GSTs的纯化及生化毒理学特性解析
采用Glutathionc Sepharose 4B亲和层析法纯化了嗜虫书虱4个地理种群(河南开封、四川广汉、重庆北碚和湖北武汉)的GSTs。结果表明,嗜虫书虱河南开封种群GSTs的纯化倍数最高,达162倍,但回收率较低,仅为29%;四川广汉种群GSTs的纯化倍数最低,仅57倍,回收率为43%;重庆北碚种群GSTs的回收率最高,达47%,纯化倍数达93倍;湖北武汉种群GSTs的回收率最低,仅为25%。应用微量滴度酶标板法的生化特性分析结果表明,河南开封种群与武汉种群GSTs对底物具有较高的催化活性,且两者之间无显著性差异(P＞0.05),但河南开封种群的GSTs对两底物表现最低的亲和力。
嗜虫书虱不同种群GSTs的最适pH研究发现,四川广汉种群的最适范围介于pH 6.5-pH 7.0之间,而其它3个种群的最适范围介于pH 7-pH 7.5之间。酶促反应最适温度研究发现,重庆北碚种群在40℃时GSTs活力最高,而其它3个种群则在35℃时GSTs的活力最高。
利尿酸、姜黄素、马来酸二乙酯和四溴磺酚钠对嗜虫书虱4个种群GSTs均有显著的离体抑制作用。综合分析发现,河南开封种群对各种抑制剂的作用最不敏感,四川广汉种群对利尿酸和马来酸二乙酯最为敏感,而重庆北碚种群对姜黄素和四溴磺酚钠最为敏感。
Psocids,belonging to Liposcelis(Psocoptera:Liposcelididae),are worldwide and commonly found in various processed and unprocessed dry foods.To date there are 123 described species in total and 27 species have been recorded in China.Outbreaks of L.bostrychophila,together with L. entoraophila could pose an alarming threat to stored product and bring about serous economic losses.In addition,the rapid development of resistance to chemical and physical treatments by the psocids has also been widely reported.
Separation and purification is the basis for protein structure and function study.The separation of a specific enzyme via protein purification methods,and the acknowledgment of the relationship between their biochemical properties and resistance to insecticides possess important significance in pest insect management.Glutathione S-transferases(GSTs,EC 2.5.1.18) are a supergene family of multifunctional enzymes found ubiquitously in mammals,insects,plants and aerobic bacteria.GSTs can metabolize foreign compounds by facilitating their reductive dehydrochlorination or by conjugation reactions with reduced glutathione,to produce water-soluble metabolites that are more readily excreted.In addition,they are involved in intracellular transport and biosynthesis of hormones,removal of toxic oxygen free radicals produced through the action of xenobiotics and protection against oxidative stress.On the basis of GSTs purification of L.bostrychophila and L. entomophila through the affinity chromatography method,the biochemical and toxicological characterizations were systematically analyzed.The study results provide biochemical and toxicological evidence for clarifying the adaptive mechanism of psocids to environment,and meanwhile enrich and develop the scientific theoretical study system for the resistance mechanism of stored product pests.The project is supported by the Program for New Century Excellent Talents in University(NCET-04-0854) and the Specialized Research Fund for the Doctoral Program of Higher Education(200806350009),and the main results are as follows:
1 Purification,biochemical and toxicological characterization of GSTs from three strains of L.bostrychophila
GSTs from three strains of L.bostrychophila(SS:susceptible strain;DDVP-R,dichlorvos-resistant strain;PH_3-R,phosphine-resistant strain) were purified through Glutathione Sepharose 4B affinity chromatography method and the purified product amounted to electrophoresis-grade purity. SDS-PAGE revealed that the molecular mass of purified GSTs was estimated at approximately 23 kDa.The N-terminus of the purified enzyme was sequenced by Edman degradation method,and a primary structure of APKYKYL TYPFNSILTGLGL was obtained.The alignment of N-terminal sequence with other insect GSTs suggested the purified protein was similar to those of Sigma class GSTs.
Purification fold of GSTs from three strains of L.bostrychophila ranged from 62 to 91 with a yield rate from 27%to 41%.The biochemical characterization result revealed that compared to SS strain,the specific activities of GSTs in DDVP-R and PH_3-R were significantly higher(P＜0.05) but no significance observed between two resistant strains(P＞0.05).Further kinetic parameters comparison showed that GSTs from DDVP-R expressed highest K_m value to the substrate CDNB while both two resistant strains exhibited lower K_m values to GSH.For V_(max),the values in resistant strains were significantly higher suggesting the catalytic ability of GSTs developed to higher levels in two resistant strains.
The pH optimum study showed that GSTs of SS strain possessed a wider optimal pH range(pH 7.0 to pH 8.0) and the optimal pH for resistant strains lied in pH 7.0 to pH 7.5.Temperature optimum of enzymatic reaction revealed that GSTs from SS and PH_3-R strains possessed highest specific activities in 35℃while activity peak recorded at 40℃for GSTs of DDVP-R strain.
In vitro inhibition of insecticides showed that carbosulfan expressed significant inhibitory effects on GSTs in three strains of L.bostrychophila and the sensitivity of GSTs from PH_3-R was lowest.For beta-cypermethrin and chlorpyrifos,the inhibition effects were not obvious and some facilitated effects appeared at lower concentrations of chlorpyrifos.Ethacrynic acid,curcumin and bromosulfalein all exhibited significant in vitro inhibition effects.Among these inhibitors,GSTs were most sensitive to curcumin with its I_(50) values at nanomolar concentrations.The comparison analysis revealed that compared to the counterparts in resistant strains GSTs from SS strain were most sensitive and sensitivity difference to inhibitors existed for GSTs in resistant strains.
2 Purification,biochemical and toxicological characterization of GSTs from different developmental stages of L.bostrychophila
GSTs from different developmental stages of L.bostrychophila(Early nymph:1st and 2nd nymph;Late nymph:3rd and 4th nymph;Adult) were purified through Glutathione Sepharose 4B affinity chromatography method.The result revealed that the purification fold of GSTs from early nymph was highest(102) with a low yield rate of 28%,while late nymph exhibited lowest purification fold(63) with a yield rate of 61%.The specific activity comparison showed that highest value observed in adults.Further kinetic parameters comparison found GSTs from early nymph expressed lowest K_m values to the substrates CDNB and GSH while adults possessed highest V_(max) values,suggesting GSTs with higher catalytic ability to the substrates existed in adults.
In vitro inhibition of chlorpyrifos showed that the pesticide expressed some inhibitory effects. At lower assay concentrations of chlorpyrifos some facilitated effects occurred while at higher concentrations inhibition appeared.The highest inhibition rate reached 39%under the test concentrations of chlorpyrifos.All five chemicals(ethacrynic acid,curcumin,disulfiram,diethyl maleate and bromosulfalein) exhibited significant in vitro inhibition on GSTs from different developmental stages of L.bostrychophila.Among these inhibitors,GSTs were least sensitive to disulfiram with its I_(50) values at millimolar concentrations.The comparison analysis revealed that GSTs in adults were most tolerant to all other inhibitors except that GSTs in late nymph least sensitive to ethacrynic acid.
Besides,the effects of metal ions(Zn~(2+),Hg~(2+),Mn~(2+),Cu~(2+) and Ca~(2+)) on GSTs of different stages of L.bostrychophila were assayed.The results suggested that all metal ions expressed significant inhibitory effects.Among these metal ions,Ca~(2+) exhibited least insensitivity with its I_(50) value at nanomolar levels.The effects of Hg~(2+) were most significant but no significant difference among different stages of L.bostrychophila(P＞0.05).
3 Purification,biochemical and toxicological characterization of GSTs from different populations of L.bostrychophila
GSTs from different populations of L.bostrychophila(Guanghan population:from Guanghan of Sichuan province;Jianyang population:from Jiangyang of Sichuan province;Lab conservation population) were purified through Glutathione Sepharose 4B affinity chromatography method.The result revealed that the purification fold of GSTs from Lab population was highest(121) with a highest yield rate of 52%,while Jianyang population exhibited lowest purification fold(40) with a lowest yield rate of 16%.The purification parameters of Guanghan population ranged between those of the other populations.Both the total protein and specific activity comparison showed that highest values recorded in Lab population but no significance occurred between the parameters of other two populations.Further kinetic parameters comparison found GSTs of Lab population expressed lowest affinity ability to the substrates CDNB and GSH,Jianyang population possessed highest affinity ability to GSH and Guanghan population showed highest affinity ability to CDNB.For V_(max) values, Lab population revealed highest catalytic activity to both substrates,significantly higher than Jianyang and Guanghan populations(P＜0.05).
In vitro inhibition of insecticides on GSTs of different populations of L.bostrychophila showed that Guanghan population was most sensitive to carbosulfan and Lab population possessed least sensitivity to the insecticide.For beta-cypermethrin,the inhibition effects were not obvious and the situation in different populations was similar.Ethacrynic acid,curcumin and diethyl maleate all exhibited significant in vitro inhibition.Among these populations,Guanghan population expressed most sensitivity to all inhibitors.
Besides,the effects of metal ions(Zn~(2+),Hg~(2+),Mn~(2+),Cu~(2+) and Ca~(2+)) on GSTs of different populations of L.bostrychophila were assayed.The results suggested that all metal ions expressed significant inhibitory effects.Among these metal ions,Ca~(2+) exhibited least insensitivity with no significant difference among different populations(P＞0.05).Similar situation observed for Mn~(2+) and Cu~(2+),Guanghan population was most sensitive to these ions while Lab population was least sensitive.
4 Purification,biochemical and toxicological characterization of GSTs from different populations of L.entomophila
GSTs from different populations of L.entomophiIa(Kaifeng population:from Kaifeng of Henan province;Guanghan population:from Guanghan of Sichuan province;Lab population:from Beibei of Chongqing city;Wuhan population:from Wuhan of Hubei province) were purified through Glutathione Sepharose 4B affinity chromatography method.The result revealed that the purification fold of GSTs from Kaifeng population was highest(162) with a yield rate of 29%,Guanghan population exhibited lowest purification fold(57) with a yield rate of 43%,Lab population possessed highest yield rate of 47%with a purification fold of 93 and the lowest yield rate was observed in Wuhan population.The activity assay showed that Kaifeng and Wuhan populations expressed relatively higher catalytic activity(P＞0.05),no significant difference between these two populations but Kaifeng population exhibited lowest affinity ability to the substrates GSH and CDNB.
The pH optimum study showed that the optimal pH for GSTs of Guanghan population of L. entomophila ranged from pH 6.5 to pH 7.0 while the range for other three populations lies in pH 7.0 to pH 7.5.Temperature optimum of enzymatic reaction revealed that GSTs from Lab population possessed highest specific activity in 40℃while activity peak recorded at 35℃for GSTs of other populations.
Ethacrynic acid,curcumin,diethyl maleate and bromosulfalein all exhibited significant in vitro inhibition effects.The comparison analysis revealed that GSTs of Kaifeng population were least sensitive to all test inhibitors,Guanghan population exhibited most sensitivity to ethacrynic acid and diethyl maleate while Lab population expressed most sensitivity to curcumin and bromosulfalein.
蛋白质分离和纯化是进行蛋白质结构和功能研究的基础。采用蛋白质纯化的方法分离参与抗药性产生的酶,明确其生物化学特性及其与抗药性的关系,对于害虫抗药性治理具有重要的意义。谷胱甘肽S-转移酶(Glutathione S-transferases,GSTs,EC 2.5.1.18)是一类多功能的超基因家族酶,广泛分布于哺乳动物、昆虫、植物以及微生物等生物体内。这类酶能催化内源还原性谷胱甘肽(GSH)与各种有害的亲电性底物相结合,增加后者的可溶性从而有利于其从细胞内排出,进而保护生物体内的核酸和蛋白质免受亲电基团攻击。同时,它也参与激素的胞内运输、合成,清除外源化合物所产生的有害的氧自由基以及保护细胞免遭氧化压力胁迫。本学位论文在教育部新世纪优秀人才支持计划(NCET-04-0854)和教育部高等学校博士点专项基金(200806350009)的资助下,利用亲和层析法对嗜卷书虱和嗜虫书虱的GSTs进行分离纯化的基础上,对纯化产物的生化毒理学特性进行了较为全面的解析,研究结果为揭示GSTs介导的书虱适应环境胁迫的机理提供了生化毒理学依据,同时丰富和发展了储藏物害虫抗性机理研究的科学理论体系。主要研究结果总结如下:
1嗜卷书虱不同品系GSTs的纯化及生化毒理学特性解析
采用Glutathione Sepharose 4B亲和层析法纯化了嗜卷书虱3个品系(SS,敏感品系;DDVP-R,敌敌畏抗性品系;PH_3-R,磷化氢抗性品系)的GSTs,纯化产物达电泳级纯度。经SDS-PAGE电泳检测,纯化后GSTs的蛋白分子量约为23 kDa。利用Edman降解法对纯化产物N端前15个氨基酸测序获得的短肽为:APKYKYL TYPFNSILTGLGL,该序列与Sigma类昆虫GSTs N端氨基酸序列高度一致,表明纯化获得的GSTs属于昆虫Sigma类。
嗜卷书虱3个品系GSTs的纯化倍数在62-91倍之间,回收率介于27-41%。利用微量滴度酶标板法对纯化后的GSTs的生化特性分析表明,与SS品系相比,DDVP-R和PH_3-R品系的GSTs活性显著提高(P＜0.05),而两抗性品系间相比较差异未达显著水平(P＞0.05)。进一步的动力学比较研究发现,以CDNB为底物时,DDVP-R品系的K_m值最高;而以GSH为底物时,两抗性品系K_m值较低,说明DDVP和PH_3的胁迫选择造成GSTs对内源性化合物GSH获得了较高的亲和能力;对V_(max)而言,两抗性品系的V_(max)值均高于敏感品系,说明其催化能力提高。
嗜卷书虱3个品系GSTs活力随pH值变化的趋势大体相同,其中SS品系GSTs的最适pH范围较广,介于pH7.0-pH8.0之间,而两抗性品系的最适范围介于pH 7.0-pH 7.5之间。酶促反应最适温度研究发现,SS和PH_3-R品系在35℃时GSTs活力最高,而DDVP-R品系则在40℃时GSTs的活力最高。
杀虫剂对嗜卷书虱3个品系GSTs的离体作用研究表明,供试的杀虫剂中丁硫克百威对嗜卷书虱3个品系的GSTs的抑制作用明显,3品系间相比较PH_3-R品系GSTs对丁硫克百威的抑制作用敏感度低,抑制中浓度I_(50)分别为SS和DDVP-R品系的3.9和4.7倍。毒死蜱和高效氯氰菊酯对嗜卷书虱的GSTs仅有部分的抑制作用,毒死蜱在低浓度时还表现为一定的激活作用。利尿酸、姜黄素和四溴磺酚钠对嗜卷书虱纯化后的GSTs均有显著的离体抑制作用。其中,姜黄素的抑制作用最强,其I_(50)达纳摩尔数量级。综合分析,SS品系与两抗性品系相比对上述抑制剂更为敏感,两抗性品系GSTs对各抑制剂的敏感性存在一定的差异。
2嗜卷书虱不同发育阶段GSTs的纯化及生化毒理学特性解析
采用Glutathione Sepharose 4B亲和层析法纯化了嗜卷书虱3个发育阶段(1-2龄低龄若虫、3-4龄高龄若虫和成虫)的GSTs。结果表明,低龄若虫GSTs的纯化倍数最高,达102倍,但回收率仅为28%;高龄若虫GSTs的纯化倍数最低,仅为63倍,但回收率高达61%;成虫GSTs的纯化倍数介于低龄和高龄若虫之间,其GSTs的比活力最高,且显著高于低龄若虫(P＜0.05)。以GSH和CDNB为底物的动力学特性分析发现,低龄若虫GSTs的K_m值最低,而成虫GSTs对两底物均表现最高的催化反应速度V_(max),表明成虫期GSTs对底物的催化能力显著高于若虫期。
毒死蜱对嗜卷书虱不发育阶段GSTs纯化后的离体作用结果表明,该药剂对GSTs有一定的抑制作用,但在低浓度时表现为一定的激活作用,随着浓度的升高,抑制作用逐渐显现。在测试的5个浓度下,毒死蜱对GSTs的最高抑制率仅为39%。5种典型化合物(利尿酸、姜黄素、双硫仑、马来酸二乙酯和四溴磺酚钠)对嗜卷书虱各发育阶段GSTs均有显著的离体抑制作用,其中GSTs对双硫仑的耐受性最强,该抑制剂的离体I_(50)达毫摩尔数量级。综合分析发现,除高龄若虫对利尿酸最不敏感外,嗜卷书虱成虫对其它抑制剂均表现为最不敏感。
此外,研究了金属离子(Zn~(2+)、Hg~(2+)、Mn~(2+)、Cu~(2+)和Ca~(2+))对嗜卷书虱GSTs活性的影响。结果表明,5种金属离子对嗜卷书虱GSTs均有显著的抑制作用。嗜卷书虱各发育阶段的GSTs对Ca~(2+)最不敏感,I_(50)达纳摩尔数量级;Hg~(2+)对嗜卷书虱GSTs的离体抑制作用最为显著,但各虫态间I_(50)值无显著性差异(P＞0.05)。
3嗜卷书虱不同地理种群GSTs的纯化及生化毒理学特性解析
采用Glutathione Sepharose 4B亲和层析法纯化了嗜卷书虱3个地理种群(四川广汉、四川简阳和实验室保存种群)的GSTs。结果表明,经亲和层析后,实验室种群GSTs的纯化倍数高达121倍,回收率亦最高,达52%;四川广汉种群纯化倍数和回收率分别为58和33%;简阳种群的纯化倍数最低,仅为40倍,回收率仅为16%。总蛋白和活力测定结果显示,实验室种群GSTs纯化产物的总蛋白含量和酶活力显著高于其余两个种群(P＜0.05),但其它种群之间各参数均无显著性差异(P＞0.05)。
分别以CDNB和GSH为底物,利用双倒数作图法获得了嗜卷书虱3个种群GSTs纯化后的动力学参数。比较分析K_m值发现,实验室种群对2种底物的亲和力最低,简阳种群对底物GSH的亲和力最高,而广汉种群对底物CDNB的亲和力最高;就V_(max)而言,实验室种群对底物GSH和CDNB均表现出最高的催化活性,且显著高于简阳种群与广汉种群(P＜0.05)。
杀虫剂的离体抑制作用测定结果表明,广汉种群对丁硫克百威最为敏感,简阳种群其次,实验室种群对该药剂的离体抑制作用最不敏感。高效氯氰菊酯对嗜卷书虱不同种群纯化后的GSTs有部分的抑制作用,且各种群受到的离体作用类似。利尿酸、姜黄素、马来酸二乙酯和四溴磺酚钠对嗜卷书虱3个种群均表现为显著的抑制作用,其中广汉种群GSTs对各种抑制剂的作用最为敏感。
各金属离子对嗜卷书虱3个种群的作用结果显示,GSTs对Ca~(2+)最不敏感,其中实验室种群与其它两种群相比更不敏感,其I_(50)值最高;Hg~(2+)对GSTs离体抑制作用最为强烈,各种群间I_(50)值无显著性差异(P＞0.05);Mn~(2+)和Ca~(2+)对各种群GSTs的离体作用较为相似,其中广汉种群对这2种金属离子的抑制作用最为敏感,而实验室种群最不敏感。
4嗜虫书虱不同地理种群GSTs的纯化及生化毒理学特性解析
采用Glutathionc Sepharose 4B亲和层析法纯化了嗜虫书虱4个地理种群(河南开封、四川广汉、重庆北碚和湖北武汉)的GSTs。结果表明,嗜虫书虱河南开封种群GSTs的纯化倍数最高,达162倍,但回收率较低,仅为29%;四川广汉种群GSTs的纯化倍数最低,仅57倍,回收率为43%;重庆北碚种群GSTs的回收率最高,达47%,纯化倍数达93倍;湖北武汉种群GSTs的回收率最低,仅为25%。应用微量滴度酶标板法的生化特性分析结果表明,河南开封种群与武汉种群GSTs对底物具有较高的催化活性,且两者之间无显著性差异(P＞0.05),但河南开封种群的GSTs对两底物表现最低的亲和力。
嗜虫书虱不同种群GSTs的最适pH研究发现,四川广汉种群的最适范围介于pH 6.5-pH 7.0之间,而其它3个种群的最适范围介于pH 7-pH 7.5之间。酶促反应最适温度研究发现,重庆北碚种群在40℃时GSTs活力最高,而其它3个种群则在35℃时GSTs的活力最高。
利尿酸、姜黄素、马来酸二乙酯和四溴磺酚钠对嗜虫书虱4个种群GSTs均有显著的离体抑制作用。综合分析发现,河南开封种群对各种抑制剂的作用最不敏感,四川广汉种群对利尿酸和马来酸二乙酯最为敏感,而重庆北碚种群对姜黄素和四溴磺酚钠最为敏感。
Psocids,belonging to Liposcelis(Psocoptera:Liposcelididae),are worldwide and commonly found in various processed and unprocessed dry foods.To date there are 123 described species in total and 27 species have been recorded in China.Outbreaks of L.bostrychophila,together with L. entoraophila could pose an alarming threat to stored product and bring about serous economic losses.In addition,the rapid development of resistance to chemical and physical treatments by the psocids has also been widely reported.
Separation and purification is the basis for protein structure and function study.The separation of a specific enzyme via protein purification methods,and the acknowledgment of the relationship between their biochemical properties and resistance to insecticides possess important significance in pest insect management.Glutathione S-transferases(GSTs,EC 2.5.1.18) are a supergene family of multifunctional enzymes found ubiquitously in mammals,insects,plants and aerobic bacteria.GSTs can metabolize foreign compounds by facilitating their reductive dehydrochlorination or by conjugation reactions with reduced glutathione,to produce water-soluble metabolites that are more readily excreted.In addition,they are involved in intracellular transport and biosynthesis of hormones,removal of toxic oxygen free radicals produced through the action of xenobiotics and protection against oxidative stress.On the basis of GSTs purification of L.bostrychophila and L. entomophila through the affinity chromatography method,the biochemical and toxicological characterizations were systematically analyzed.The study results provide biochemical and toxicological evidence for clarifying the adaptive mechanism of psocids to environment,and meanwhile enrich and develop the scientific theoretical study system for the resistance mechanism of stored product pests.The project is supported by the Program for New Century Excellent Talents in University(NCET-04-0854) and the Specialized Research Fund for the Doctoral Program of Higher Education(200806350009),and the main results are as follows:
1 Purification,biochemical and toxicological characterization of GSTs from three strains of L.bostrychophila
GSTs from three strains of L.bostrychophila(SS:susceptible strain;DDVP-R,dichlorvos-resistant strain;PH_3-R,phosphine-resistant strain) were purified through Glutathione Sepharose 4B affinity chromatography method and the purified product amounted to electrophoresis-grade purity. SDS-PAGE revealed that the molecular mass of purified GSTs was estimated at approximately 23 kDa.The N-terminus of the purified enzyme was sequenced by Edman degradation method,and a primary structure of APKYKYL TYPFNSILTGLGL was obtained.The alignment of N-terminal sequence with other insect GSTs suggested the purified protein was similar to those of Sigma class GSTs.
Purification fold of GSTs from three strains of L.bostrychophila ranged from 62 to 91 with a yield rate from 27%to 41%.The biochemical characterization result revealed that compared to SS strain,the specific activities of GSTs in DDVP-R and PH_3-R were significantly higher(P＜0.05) but no significance observed between two resistant strains(P＞0.05).Further kinetic parameters comparison showed that GSTs from DDVP-R expressed highest K_m value to the substrate CDNB while both two resistant strains exhibited lower K_m values to GSH.For V_(max),the values in resistant strains were significantly higher suggesting the catalytic ability of GSTs developed to higher levels in two resistant strains.
The pH optimum study showed that GSTs of SS strain possessed a wider optimal pH range(pH 7.0 to pH 8.0) and the optimal pH for resistant strains lied in pH 7.0 to pH 7.5.Temperature optimum of enzymatic reaction revealed that GSTs from SS and PH_3-R strains possessed highest specific activities in 35℃while activity peak recorded at 40℃for GSTs of DDVP-R strain.
In vitro inhibition of insecticides showed that carbosulfan expressed significant inhibitory effects on GSTs in three strains of L.bostrychophila and the sensitivity of GSTs from PH_3-R was lowest.For beta-cypermethrin and chlorpyrifos,the inhibition effects were not obvious and some facilitated effects appeared at lower concentrations of chlorpyrifos.Ethacrynic acid,curcumin and bromosulfalein all exhibited significant in vitro inhibition effects.Among these inhibitors,GSTs were most sensitive to curcumin with its I_(50) values at nanomolar concentrations.The comparison analysis revealed that compared to the counterparts in resistant strains GSTs from SS strain were most sensitive and sensitivity difference to inhibitors existed for GSTs in resistant strains.
2 Purification,biochemical and toxicological characterization of GSTs from different developmental stages of L.bostrychophila
GSTs from different developmental stages of L.bostrychophila(Early nymph:1st and 2nd nymph;Late nymph:3rd and 4th nymph;Adult) were purified through Glutathione Sepharose 4B affinity chromatography method.The result revealed that the purification fold of GSTs from early nymph was highest(102) with a low yield rate of 28%,while late nymph exhibited lowest purification fold(63) with a yield rate of 61%.The specific activity comparison showed that highest value observed in adults.Further kinetic parameters comparison found GSTs from early nymph expressed lowest K_m values to the substrates CDNB and GSH while adults possessed highest V_(max) values,suggesting GSTs with higher catalytic ability to the substrates existed in adults.
In vitro inhibition of chlorpyrifos showed that the pesticide expressed some inhibitory effects. At lower assay concentrations of chlorpyrifos some facilitated effects occurred while at higher concentrations inhibition appeared.The highest inhibition rate reached 39%under the test concentrations of chlorpyrifos.All five chemicals(ethacrynic acid,curcumin,disulfiram,diethyl maleate and bromosulfalein) exhibited significant in vitro inhibition on GSTs from different developmental stages of L.bostrychophila.Among these inhibitors,GSTs were least sensitive to disulfiram with its I_(50) values at millimolar concentrations.The comparison analysis revealed that GSTs in adults were most tolerant to all other inhibitors except that GSTs in late nymph least sensitive to ethacrynic acid.
Besides,the effects of metal ions(Zn~(2+),Hg~(2+),Mn~(2+),Cu~(2+) and Ca~(2+)) on GSTs of different stages of L.bostrychophila were assayed.The results suggested that all metal ions expressed significant inhibitory effects.Among these metal ions,Ca~(2+) exhibited least insensitivity with its I_(50) value at nanomolar levels.The effects of Hg~(2+) were most significant but no significant difference among different stages of L.bostrychophila(P＞0.05).
3 Purification,biochemical and toxicological characterization of GSTs from different populations of L.bostrychophila
GSTs from different populations of L.bostrychophila(Guanghan population:from Guanghan of Sichuan province;Jianyang population:from Jiangyang of Sichuan province;Lab conservation population) were purified through Glutathione Sepharose 4B affinity chromatography method.The result revealed that the purification fold of GSTs from Lab population was highest(121) with a highest yield rate of 52%,while Jianyang population exhibited lowest purification fold(40) with a lowest yield rate of 16%.The purification parameters of Guanghan population ranged between those of the other populations.Both the total protein and specific activity comparison showed that highest values recorded in Lab population but no significance occurred between the parameters of other two populations.Further kinetic parameters comparison found GSTs of Lab population expressed lowest affinity ability to the substrates CDNB and GSH,Jianyang population possessed highest affinity ability to GSH and Guanghan population showed highest affinity ability to CDNB.For V_(max) values, Lab population revealed highest catalytic activity to both substrates,significantly higher than Jianyang and Guanghan populations(P＜0.05).
In vitro inhibition of insecticides on GSTs of different populations of L.bostrychophila showed that Guanghan population was most sensitive to carbosulfan and Lab population possessed least sensitivity to the insecticide.For beta-cypermethrin,the inhibition effects were not obvious and the situation in different populations was similar.Ethacrynic acid,curcumin and diethyl maleate all exhibited significant in vitro inhibition.Among these populations,Guanghan population expressed most sensitivity to all inhibitors.
Besides,the effects of metal ions(Zn~(2+),Hg~(2+),Mn~(2+),Cu~(2+) and Ca~(2+)) on GSTs of different populations of L.bostrychophila were assayed.The results suggested that all metal ions expressed significant inhibitory effects.Among these metal ions,Ca~(2+) exhibited least insensitivity with no significant difference among different populations(P＞0.05).Similar situation observed for Mn~(2+) and Cu~(2+),Guanghan population was most sensitive to these ions while Lab population was least sensitive.
4 Purification,biochemical and toxicological characterization of GSTs from different populations of L.entomophila
GSTs from different populations of L.entomophiIa(Kaifeng population:from Kaifeng of Henan province;Guanghan population:from Guanghan of Sichuan province;Lab population:from Beibei of Chongqing city;Wuhan population:from Wuhan of Hubei province) were purified through Glutathione Sepharose 4B affinity chromatography method.The result revealed that the purification fold of GSTs from Kaifeng population was highest(162) with a yield rate of 29%,Guanghan population exhibited lowest purification fold(57) with a yield rate of 43%,Lab population possessed highest yield rate of 47%with a purification fold of 93 and the lowest yield rate was observed in Wuhan population.The activity assay showed that Kaifeng and Wuhan populations expressed relatively higher catalytic activity(P＞0.05),no significant difference between these two populations but Kaifeng population exhibited lowest affinity ability to the substrates GSH and CDNB.
The pH optimum study showed that the optimal pH for GSTs of Guanghan population of L. entomophila ranged from pH 6.5 to pH 7.0 while the range for other three populations lies in pH 7.0 to pH 7.5.Temperature optimum of enzymatic reaction revealed that GSTs from Lab population possessed highest specific activity in 40℃while activity peak recorded at 35℃for GSTs of other populations.
Ethacrynic acid,curcumin,diethyl maleate and bromosulfalein all exhibited significant in vitro inhibition effects.The comparison analysis revealed that GSTs of Kaifeng population were least sensitive to all test inhibitors,Guanghan population exhibited most sensitivity to ethacrynic acid and diethyl maleate while Lab population expressed most sensitivity to curcumin and bromosulfalein.
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