基于外来生物B型烟粉虱重要唾液组分的入侵机理研究
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摘要
重大入侵生物B型烟粉虱自20世纪末传入我国以来，对我国的生态系统和农业生产造成了严重危害，极强的竞争能力和极高的寄主适应性是其暴发成灾的主要原因。当B型烟粉虱传入到一个新的生境，通过口针从寄主植物持续地获取营养物质，是其完成世代发育和种群扩增的必备因素；与此同时，B型烟粉虱还通过口针将其唾液注入到植物组织，实现两者之间的液体物质交流，这种交流是入侵粉虱对本地生物群落产生作用最主要的方式之一。外来物种的入侵机制是入侵生物学的核心问题，各种入侵假说已成为生物入侵研究的重要理论和蓝本。“内在优势假说”阐明了成功入侵者相对于本地竞争者的潜在优势，“新式武器假说”则强调了入侵者对本地生物群落的化感作用。因此，B型烟粉虱是否可以利用唾液作为武器影响本地生物群落，而这种影响是否为本地近缘种温室粉虱所不具备的竞争优势，成为了本研究最核心的科学问题。
本文首先通过比较不同饲养时间，密度效应，饲料蔗糖浓度和饲料pH值对B型烟粉虱和温室粉虱死亡率的影响，优化了粉虱的人工饲养体系，结果表明采用400头粉虱，饲料蔗糖浓度10%，pH值6.5，饲养两天后能够确保大部分粉虱稳定取食。随后采用人工饲养结合生物化学的方法，对两种粉虱的唾液水解酶、氧化酶和果胶酶进行了鉴定和分析。对两种粉虱唾液水解酶的分析表明，其在各自的特异嗜好寄主上都有更高的唾液碱性磷酸酶比活力，同时B型烟粉虱番茄和棉花种群的唾液碱性磷酸酶比活力要显著大于温室粉虱对应种群的酶比活力。两种粉虱唾液碱性磷酸酶活性随着饲料蔗糖浓度的降低而升高，表明其可能帮助粉虱代谢植物中的蔗糖。对两种粉虱唾液氧化酶的分析表明，B型烟粉虱唾液多酚氧化酶与过氧化物酶的比活力显著高于温室粉虱对应酶的比活力，两种粉虱唾液多酚氧化酶均能与五种酚类底物反应，特异性反应顺序为：五倍子酸>栎精>邻苯二酚>咖啡酸>绿原酸，B型烟粉虱唾液多酚氧化酶对不同酚类底物的降解能力均显著高于温室粉虱。致病疫霉菌侵染、机械损伤、B型烟粉虱预取食、外施水杨酸四种诱导抗性处理能在不同程度上诱导影响两种粉虱唾液多酚氧化酶的比活力上升，这四种处理后B型烟粉虱唾液多酚氧化酶的比活力变化均显著高于温室粉虱。同时B型烟粉虱取食能够使番茄叶片的总酚含量升高，这可能对两种粉虱具有偏利效应。本研究未能在两种粉虱唾液中鉴定出果胶甲酯酶和多聚半乳糖醛酸酶。
克隆了B型烟粉虱多酚氧化物家族漆酶-1基因，序列分析表明B型烟粉虱唾液腺很可能将水溶且高活性的漆酶-1分泌到唾液中，一方面促进胶状唾液的快速氧化而形成口针鞘，另一方面降解植物组织中特定的酚类物质而帮助取食，而这些功能很可能在刺吸式昆虫漆酶-1基因中高度保守。总的来说，B型烟粉虱利用碱性磷酸酶和多酚氧化酶适应寄主的能力要强于温室粉虱，这可能是其能够成功入侵的内在优势之一。
Bemisia tabaci (Gennadius) biotype B was introduced to China at the end of20th century andposed huge threat to the ecosystem and agricultural industry. Strong competitiveness and high hostadaptation were the main causes for its outbreak. B.tabaci biotype B is piercing-sucking insect, whichwill secrete gell saliva and watery saliva when it sucks phloem sap from the plants. This liquidsexchange between whitefly and plant is one of the most important ways how invasive whitefly affectsthe local biota. Invasion hypothesis are the key theories of invasion biology. Inherent superiorityhypothesis describes the potential advantages of successful invader, while novel weapon hypothesisemphasis the allelopathic effect of invader on the local community. Thus the key scientific questions inthis study are: Is saliva a novel weapon for B.tabaci biotype B? Can this be the Inherent superiority forB.tabaci biotype B when compare to the indigenous species Trialeurodes vaporariorum?
In this study, we first tested the effects of rearing time, density, sucrose concentration of diet andpH value of diet on the mortality of the B.tabaci biotype B and T. vaporariorum. The results showedrearing400whiteflies on10%sucrose concentration and pH6.5diet for2days is the best combinationfor saliva collection. This artificial feeding system and biochemical methods were used to identify andanalyze different enzymes in whitefly saliva. For salivary hydrolase, both B.tabaci biotype B and T.vaporariorum have higher salivary alkaline phosphatase activities on their preferred hosts compared tothose from unsuitable hosts. B. tabaci B-biotype maintained higher salivary alkaline phosphataseactivity than T. vaporariorum on tomato and cotton. The salivary alkaline phosphatase activities fromboth whiteflies increased when the sucrose concentration in diet decrease, which suggest alkalinephosphatase plays a role in sucrose metabolism. For salivary pectinase, Pectin methylesterase andpolygalacturonase were not identified in the saliva of two whitefly species in this study.
For salivary oxides, polyphenol oxidase and peroxidase were found in the gel and watery saliva ofboth B. tabaci B-biotype and T.vaporariorum. For tomato colonies, the polyphenol oxidase andperoxidase activities in the watery saliva of B. tabaci B-biotype were2.27and1.34fold higher thanthose of T. vaporariorum. The polyphenol oxidase activities against specific phenolic compoundscommonly found in plants were compared. while the order of activity remained the same for bothwhitefly species, gallic acid>quercetin> pyrocatechol> caffeic acid>chlorogenic acid, the level ofactivity was markedly different with B. tabaci B-biotype consistently demonstrating much higher levelsof activity We also measured polyphenol oxidase activity in both species after they had feed on plantsthat were undamaged or had been previously damaged with either a plant pathogen (Phytophthorainfestans) infection, mechanical damage, B. tabaci B-biotype infestation or exogenous salicylic acid.For B. tabaci B-biotype, polyphenol oxidase activities in watery saliva increased229%,184%,152%,and139%in response to the four treatments while those of T. vaporariorum only increased133%,119%,113%,103%, respectively. B. tabaci B-biotype infestation can significantly increase the total phenolicscontent of tomato leaves. Meanwhile, feeding on tomato pre-infested with B. tabaci B-biotype had no significant effect on the survival rate of B. tabaci B-biotype but can decrease the survival rate of T.vaporariorum significantly.
Laccase belong to the polyphenol oxidase family and is known to be present in the saliva andsalivary gland of several insect species. The full-length cDNA sequence of Laccase-1gene(BtLac-1)was cloned from B.tabaci biotype B, which was sequenced to be3261bp and encoded911amino acid.BtLac-1has the typical features of Cu-oxidase domain, which belongs to blue copper-containingpolyphenol oxidases family. Homology analysis of amino acid sequence showed that BtLac-1shares58%identity with Nephotettix cincticeps and Acyrthosiphon pisum. The sequence analyses suggest thatB.tabaci biotype B produce soluble laccase in an active form. This indicates that the salivary laccasecauses rapid oxidation of phenolic substances just after being released into the saliva. In general, theseresults suggest that B. tabaci B-biotype is better adapted with the greater alkaline phosphatase andpolyphenol oxidase activities to metabolize sucrose or detoxify high concentrations of phenolics than T.vaporariorum, and have a significant advantage to help itself hold high fitness on plants, resulting rapidand successful invasion.
本文首先通过比较不同饲养时间，密度效应，饲料蔗糖浓度和饲料pH值对B型烟粉虱和温室粉虱死亡率的影响，优化了粉虱的人工饲养体系，结果表明采用400头粉虱，饲料蔗糖浓度10%，pH值6.5，饲养两天后能够确保大部分粉虱稳定取食。随后采用人工饲养结合生物化学的方法，对两种粉虱的唾液水解酶、氧化酶和果胶酶进行了鉴定和分析。对两种粉虱唾液水解酶的分析表明，其在各自的特异嗜好寄主上都有更高的唾液碱性磷酸酶比活力，同时B型烟粉虱番茄和棉花种群的唾液碱性磷酸酶比活力要显著大于温室粉虱对应种群的酶比活力。两种粉虱唾液碱性磷酸酶活性随着饲料蔗糖浓度的降低而升高，表明其可能帮助粉虱代谢植物中的蔗糖。对两种粉虱唾液氧化酶的分析表明，B型烟粉虱唾液多酚氧化酶与过氧化物酶的比活力显著高于温室粉虱对应酶的比活力，两种粉虱唾液多酚氧化酶均能与五种酚类底物反应，特异性反应顺序为：五倍子酸>栎精>邻苯二酚>咖啡酸>绿原酸，B型烟粉虱唾液多酚氧化酶对不同酚类底物的降解能力均显著高于温室粉虱。致病疫霉菌侵染、机械损伤、B型烟粉虱预取食、外施水杨酸四种诱导抗性处理能在不同程度上诱导影响两种粉虱唾液多酚氧化酶的比活力上升，这四种处理后B型烟粉虱唾液多酚氧化酶的比活力变化均显著高于温室粉虱。同时B型烟粉虱取食能够使番茄叶片的总酚含量升高，这可能对两种粉虱具有偏利效应。本研究未能在两种粉虱唾液中鉴定出果胶甲酯酶和多聚半乳糖醛酸酶。
克隆了B型烟粉虱多酚氧化物家族漆酶-1基因，序列分析表明B型烟粉虱唾液腺很可能将水溶且高活性的漆酶-1分泌到唾液中，一方面促进胶状唾液的快速氧化而形成口针鞘，另一方面降解植物组织中特定的酚类物质而帮助取食，而这些功能很可能在刺吸式昆虫漆酶-1基因中高度保守。总的来说，B型烟粉虱利用碱性磷酸酶和多酚氧化酶适应寄主的能力要强于温室粉虱，这可能是其能够成功入侵的内在优势之一。
Bemisia tabaci (Gennadius) biotype B was introduced to China at the end of20th century andposed huge threat to the ecosystem and agricultural industry. Strong competitiveness and high hostadaptation were the main causes for its outbreak. B.tabaci biotype B is piercing-sucking insect, whichwill secrete gell saliva and watery saliva when it sucks phloem sap from the plants. This liquidsexchange between whitefly and plant is one of the most important ways how invasive whitefly affectsthe local biota. Invasion hypothesis are the key theories of invasion biology. Inherent superiorityhypothesis describes the potential advantages of successful invader, while novel weapon hypothesisemphasis the allelopathic effect of invader on the local community. Thus the key scientific questions inthis study are: Is saliva a novel weapon for B.tabaci biotype B? Can this be the Inherent superiority forB.tabaci biotype B when compare to the indigenous species Trialeurodes vaporariorum?
In this study, we first tested the effects of rearing time, density, sucrose concentration of diet andpH value of diet on the mortality of the B.tabaci biotype B and T. vaporariorum. The results showedrearing400whiteflies on10%sucrose concentration and pH6.5diet for2days is the best combinationfor saliva collection. This artificial feeding system and biochemical methods were used to identify andanalyze different enzymes in whitefly saliva. For salivary hydrolase, both B.tabaci biotype B and T.vaporariorum have higher salivary alkaline phosphatase activities on their preferred hosts compared tothose from unsuitable hosts. B. tabaci B-biotype maintained higher salivary alkaline phosphataseactivity than T. vaporariorum on tomato and cotton. The salivary alkaline phosphatase activities fromboth whiteflies increased when the sucrose concentration in diet decrease, which suggest alkalinephosphatase plays a role in sucrose metabolism. For salivary pectinase, Pectin methylesterase andpolygalacturonase were not identified in the saliva of two whitefly species in this study.
For salivary oxides, polyphenol oxidase and peroxidase were found in the gel and watery saliva ofboth B. tabaci B-biotype and T.vaporariorum. For tomato colonies, the polyphenol oxidase andperoxidase activities in the watery saliva of B. tabaci B-biotype were2.27and1.34fold higher thanthose of T. vaporariorum. The polyphenol oxidase activities against specific phenolic compoundscommonly found in plants were compared. while the order of activity remained the same for bothwhitefly species, gallic acid>quercetin> pyrocatechol> caffeic acid>chlorogenic acid, the level ofactivity was markedly different with B. tabaci B-biotype consistently demonstrating much higher levelsof activity We also measured polyphenol oxidase activity in both species after they had feed on plantsthat were undamaged or had been previously damaged with either a plant pathogen (Phytophthorainfestans) infection, mechanical damage, B. tabaci B-biotype infestation or exogenous salicylic acid.For B. tabaci B-biotype, polyphenol oxidase activities in watery saliva increased229%,184%,152%,and139%in response to the four treatments while those of T. vaporariorum only increased133%,119%,113%,103%, respectively. B. tabaci B-biotype infestation can significantly increase the total phenolicscontent of tomato leaves. Meanwhile, feeding on tomato pre-infested with B. tabaci B-biotype had no significant effect on the survival rate of B. tabaci B-biotype but can decrease the survival rate of T.vaporariorum significantly.
Laccase belong to the polyphenol oxidase family and is known to be present in the saliva andsalivary gland of several insect species. The full-length cDNA sequence of Laccase-1gene(BtLac-1)was cloned from B.tabaci biotype B, which was sequenced to be3261bp and encoded911amino acid.BtLac-1has the typical features of Cu-oxidase domain, which belongs to blue copper-containingpolyphenol oxidases family. Homology analysis of amino acid sequence showed that BtLac-1shares58%identity with Nephotettix cincticeps and Acyrthosiphon pisum. The sequence analyses suggest thatB.tabaci biotype B produce soluble laccase in an active form. This indicates that the salivary laccasecauses rapid oxidation of phenolic substances just after being released into the saliva. In general, theseresults suggest that B. tabaci B-biotype is better adapted with the greater alkaline phosphatase andpolyphenol oxidase activities to metabolize sucrose or detoxify high concentrations of phenolics than T.vaporariorum, and have a significant advantage to help itself hold high fitness on plants, resulting rapidand successful invasion.
引文
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