B型烟粉虱取食烟草对烟蚜的影响及生理机制探讨
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摘要
B型烟粉虱自入侵我国以来,迅速扩展蔓延,已成为多种蔬菜、棉花、烟草等作物上的害虫优势种。我们在田间调查时发现烟粉虱发生的烟草植株上很少有烟蚜发生,说明B型烟粉虱在与烟蚜种间竞争中占据明显的优势,而目前有关B型烟粉虱诱导的寄主植物防御反应对同为韧皮部取食的昆虫的影响国内外均未见报道。本研究从田间现象入手,以模式植物—烟草作为寄主材料,以烟粉虱—烟草—烟蚜作为研究体系,研究了B型烟粉虱诱导的植物防御反应对烟蚜种群的抑制作用和主要影响因素,并进行了烟粉虱为害烟草对烟蚜产生防御的抗虫机制初探。本研究对明确烟粉虱对烟蚜的竞争取代能力,揭示烟粉虱对烟蚜的种间竞争取代机制,深入认识种群动态机制,有效的实施害虫可持续治理具有指导价值。本研究主要结果如下:
(1)研究了四个烟草品种对B型烟粉虱生长发育和繁殖的影响,结果表明K326、CF-965(中烟100)是易感虫品种,其上的烟粉虱发育历期较短,存活率较高,繁殖力较强;而NC-82、NC-89是较抗虫品种,其上的烟粉虱发育历期明显延长,存活率降低,繁殖力减少,不利于烟粉虱的生长发育和繁殖。
(2)采用“H”型嗅觉仪法和离体叶片选择法两种方法研究了烟粉虱为害的烟草植株和植株上不同叶位的叶片对烟蚜寄主选择的影响。结果表明烟粉虱为害后的烟草植株对烟蚜有明显的驱避作用;烟粉虱为害后的烟草不同叶位的叶片对烟蚜有不同的影响,与各自的对照叶片相比较,虫体叶对烟蚜有明显的排斥作用,而系统白脉叶和心叶则有明显的引诱作用。
(3)烟粉虱为害后的烟草植株对后侵染的烟蚜种群动态和个体生长发育及繁殖均有明显的抑制作用。烟粉虱混合虫态和若虫前侵染的烟草植株对当代后侵染的烟蚜死亡率均有显著影响,分别较对照植株升高了23.8%、30.0%;烟蚜第二代、第三代的种群增长倍数也较对照植株分别降低了55.5%、49.2%;87.9%、79.6%,烟蚜的种群增长受到显著的抑制作用。利用小叶笼法研究了烟粉虱为害烟草对烟蚜生长发育和繁殖的影响,结果表明,烟粉虱为害后的烟草系统白脉叶片对烟蚜影响显著,与无烟粉虱为害的对照叶片相比较,接烟蚜后第8天死亡率升高46.6%,发育历期延长0.9天,平均相对生长率降低6.5%,平均单雌产仔量减少5.7头/雌,生长发育和繁殖受到明显的抑制作用;但烟粉虱为害的烟草局部叶片对烟蚜没有明显的负面影响。
(4)研究了烟粉虱为害虫态、为害密度、为害时间等因素对烟粉虱诱导的烟草对烟蚜防御反应的影响。结果证明:烟粉虱若虫是诱导烟草对烟蚜防御反应的主要虫态,而500头/株烟粉虱成虫持续为害9天,没能诱导出烟草对烟蚜的防御反应。
设计了烟粉虱三龄若虫为害四个密度处理,证明烟粉虱的为害密度对烟蚜的诱导防御反应有阈值效应。烟粉虱若虫密度为0—1头/cm~2对烟蚜无不利影响;当若虫密度为2—3头/cm~2时烟草开始表现对烟蚜的防御反应;当若虫密度为5—6头/cm~2时,对烟蚜的抑制作用最明显,死亡率较对照升高56.8%;当烟粉虱若虫密度达11—12头/cm~2时,对烟蚜的负面影响较上一个密度略有降低。
在9—10头/cm~2烟粉虱密度条件下,随着烟粉虱若虫生长和为害时间的延长,对烟蚜防御反应的影响呈现先逐渐增强然后减弱的的现象。烟粉虱若虫为害1天时烟草白脉叶片对烟蚜没有明显影响;为害第5天,白脉叶片上烟蚜的死亡率较对照叶片升高36.1%;为害第10天烟草对烟蚜的防御反应表现最强烈,白脉叶上烟蚜的死亡率较对照叶片升高72.8%,平均相对生长率最低;为害第15天时烟粉虱若虫处于4龄拟蛹期,对烟蚜的影响较为害10天时减小,死亡率较对照植株升高58.9%。
(5)证明了烟粉虱取食诱导的烟草对烟蚜的防御反应具有明显的时空效应。采用小叶笼法研究了烟粉虱为害烟草对烟蚜死亡率、相对生长率和繁殖力的影响。研究结果表明,烟粉虱为害后的烟草植株四个叶位中,其中尤以白脉叶影响显著,影响顺序为白脉叶＞中间叶＞心叶＞虫体叶,证明了烟粉虱诱导的烟草对烟蚜的防御反应在烟草植株体内具有空间传递性。在烟粉虱发育至3龄时去除烟粉虱若虫虫体,然后接烟蚜,在去虫后15天内对烟蚜死亡率及相对生长率的影响与不去虫的对照相比均有明显差异,但随着去虫时间的延长,烟草对烟蚜的防御反应逐渐减弱;至去虫后第20天这种负面影响消失,说明烟粉虱取食诱导的烟草抗虫性能持续表达15天左右。
(6)测定了烟粉虱为害的烟草体内主要营养指标的变化。通过可溶性糖、可溶性蛋白含量变化与对烟蚜抑制影响的时间和空间分析,表明可溶性糖、可溶性蛋白的含量与烟蚜的死亡率呈密切的负相关性;受烟粉虱为害后的烟草白脉叶中氨基酸总含量上升,其中,天门冬氨酸含量降低,脯氨酸、丙氨酸、甘氨酸、缬氨酸、苯丙氨酸及赖氨酸等大多数种类含量上升。烟粉虱为害后的烟草体内总蛋白含量的变化亦与烟草抗蚜性有一定的相关。
Bemisia tabaci(Gennadius) B biotype spread rapidly when it invades in China.It has been a key pest of various vegetables,cotton,tobacco and so on.At the present time,B.tabaci B biotype occurs in tobacco farmlands of Shandong province widely.We find that there has a few Myzus persicae(Sulzer) on tobacco infested by B.tabaci when investigating in farm.It indicates that B.tabaci was predomination in competing with M.persicae.However,there have no report about the influence of defense responses induced by B.tabaci on other sap-feeding insects at home and abroad currently.We start at farming phenomenon,choose tobacco as test materials,choose B.tabaci--tobacco-M.persicae as research system,study the effect of tobacco's defense response induced by B.tabaci on population growth,development and reproduction of M.persicae,major effect factors.We also investigate induced resistance mechanism initially.Investigating defense responses of tobacco to M.persicae induced by B.tabaci have important academic meaning for definiting competitive replacement ability of B.tabaci to M.persicae,showing competitive replacement mechanism of B.tabaci to M.persicae,and implementing sustainable control effectively.Major results as follows:
(1) The effect of four tobacco varieties on development and reproduction of B.tabaci was studied.The results showed that K326,CF-965 were infested by B.tabaci easier.The developmental periods of B.tabaci were shorter than other tobacco varieties'.Survivorship and the number of eggs deposited by B.tabaci were higher than other tobacco varieties'. NC-82,NC-89 were more resistant to B.tabaci.The developmental periods of B.tabaci feeding on them were longer,survivorship were lower,and the fecundities were lower.They went against development and reproduction of B.tabaci.
(2) The effect of B.tabaci on host plant preference of M.persicae was studied with two different methods.The results showed that M.persicae adults liked to choose the plant without B.tabaci infested,compared to host plant infested by B.tabaci.And for leaves depart from host plant,tobacco leaves of different position had different effect on host plant preference of M.persicae.Compared to respective control leaves,damaged leaves had significant repellence on M.persicae,but,white-vein leaves and unexpanded leaves had significant attraction to M.persicae.
(3) Tobacco infested by B.tabaci had significant inhibition on population growth, development and reproduction of M.persicae.Mix developmental stages and nymphs of B.tabaci infested tobacco had significant influence on mortality of first generation of M.persicae.They were higher 23.8%,30.0%than that of M.persicae fed on the plant without B.tabaci infested.Compared to control plants,the population growth rate of second generation,third generation reduced 55.5%,49.2%;87.9%、79.6%,respectively.The population growth of M.persicae was inhibited significantly.The effect of B.tabaci damaged leaves and white-vein leaves on development and reproduction of M.persicae was studied with clip-cage.The results showed that the white-vein leaves infested by B.tabaci had significant detrimental influence.Compared to leaves of same position on control plant,the mortality increased 46.6%when switching on M.persicae for 8 days,developmental times prolonged 0.9 days,mean relative growth rate reduced 6.5%,and average fecundity decreased 5.7 per female.The development and reproduction of M.persicae was inhibited significantly. But,the damaged leaves had no significant negative influence on development and reproduction of M.persicae.
(4) Major effect factors of tobacco defense responses to M.persicae induced by B.tabaci were studied.The results indicated that B.tabaci nymphs were major factor which induced defenses responses to M.persicae.Defense responses of tobacco to M.persicae could not induced by B.tabaci infested 9 days with 500 adults per plant.
The effect of infested density of B.tabaci(0-1/cm~2,2-3/cm~2,5-6/cm~2,11-12/cm~2) on M.persicae was studied.The results showed that tobacco infested by B.tabaci nymphs with 0-1/cm~2 had no harmful effect on M.persicae.Tobacco defenses to M.persicae began to expression when density reached to 2-3/cm~2.When density reached to 5-6/cm~2,it was expressed the most significantly.The mortality was the highest,56.8%higher than control. When density reached to 11-12/cm~2,the negative effect on M.persicae was lower than prior infested density.
The influence of infested time of B.tabaci nymphs with 9-10/cm~2(1d,5d,10d,15d) on M.persicae was studied.Along with growing of B.tabaci nymphs and prolonging of infested time,the influence of defense responses to M.persicae began to strengthen and then weak. B.tabaci nymphs infested 1 day had no significant effect on M.persicae.Induced defense responses of tobacco began to express when nymphs infested 5 days,the mortality of M.persicae feeding on white-vein leaves was 36.1%higher than control plant.10 days infested by B.tabaci nymphs resulted in strong expression of defenses to M.persicae.The mortality of M.persicae fed on white-vein was the highest,72.8%higher than control plants. Mean relative growth rate was the lowest.B.tabaci nymphs reached to subnymph periods when they infested 15 days.The effect on M.persicae began to reduce compared to that of 10 days infested by B.tabaci nymphs,the mortality was 58.9%higher than contrast plant.
(5) Tobacco's defense response induced by B.tabaci to M.persicae had temporal and spatial effect.We studied the effect of defense responses on mortality,population growth rate, and fecundity of M.persicae with clip-cage.The results showed that among leaves of four position infested by B.tabaci,white-vein leaves had significant influence on development of M.persicae.The sequence of effect on mortality,development and reproduction of M.persicae was white-vein leaves>middle leaves>unexpanded leaves>damaged leaves.It proved that defense responses of tobacco to M.persicae can transmit spatially within tobacco.
We wiped off nymphs when B.tabaci nymphs developed to 3_(rd) instar and then switched on M.persicae.There had significant influence on M.persicae after wiped off B.tabaci nymphs for 15 days.Mortality and population growth rate of M.persicae had significant difference between treated and control plants.Along with duration prolonged,the influence began to retard.After 20 days,the negative influence disappeared.It suggested that the duration of induced resistance was around 15 days.
(6) We determinated major nutritional index in tobacco damaged by B.tabaci.The results indicated that there had negative correlation between soluble sugar,soluble protein content and mortality of M.persicae.Total content of amino acids in tobacco infested by B.tabaci increased.Content of Asp deduced,content of Pro,Ala,Gly,Val,Phe,Lys and so on increased. The total protein content of tobacco damaged by B.tabaci also had some relativity with tobacco's resistance to M.persicae.
(1)研究了四个烟草品种对B型烟粉虱生长发育和繁殖的影响,结果表明K326、CF-965(中烟100)是易感虫品种,其上的烟粉虱发育历期较短,存活率较高,繁殖力较强;而NC-82、NC-89是较抗虫品种,其上的烟粉虱发育历期明显延长,存活率降低,繁殖力减少,不利于烟粉虱的生长发育和繁殖。
(2)采用“H”型嗅觉仪法和离体叶片选择法两种方法研究了烟粉虱为害的烟草植株和植株上不同叶位的叶片对烟蚜寄主选择的影响。结果表明烟粉虱为害后的烟草植株对烟蚜有明显的驱避作用;烟粉虱为害后的烟草不同叶位的叶片对烟蚜有不同的影响,与各自的对照叶片相比较,虫体叶对烟蚜有明显的排斥作用,而系统白脉叶和心叶则有明显的引诱作用。
(3)烟粉虱为害后的烟草植株对后侵染的烟蚜种群动态和个体生长发育及繁殖均有明显的抑制作用。烟粉虱混合虫态和若虫前侵染的烟草植株对当代后侵染的烟蚜死亡率均有显著影响,分别较对照植株升高了23.8%、30.0%;烟蚜第二代、第三代的种群增长倍数也较对照植株分别降低了55.5%、49.2%;87.9%、79.6%,烟蚜的种群增长受到显著的抑制作用。利用小叶笼法研究了烟粉虱为害烟草对烟蚜生长发育和繁殖的影响,结果表明,烟粉虱为害后的烟草系统白脉叶片对烟蚜影响显著,与无烟粉虱为害的对照叶片相比较,接烟蚜后第8天死亡率升高46.6%,发育历期延长0.9天,平均相对生长率降低6.5%,平均单雌产仔量减少5.7头/雌,生长发育和繁殖受到明显的抑制作用;但烟粉虱为害的烟草局部叶片对烟蚜没有明显的负面影响。
(4)研究了烟粉虱为害虫态、为害密度、为害时间等因素对烟粉虱诱导的烟草对烟蚜防御反应的影响。结果证明:烟粉虱若虫是诱导烟草对烟蚜防御反应的主要虫态,而500头/株烟粉虱成虫持续为害9天,没能诱导出烟草对烟蚜的防御反应。
设计了烟粉虱三龄若虫为害四个密度处理,证明烟粉虱的为害密度对烟蚜的诱导防御反应有阈值效应。烟粉虱若虫密度为0—1头/cm~2对烟蚜无不利影响;当若虫密度为2—3头/cm~2时烟草开始表现对烟蚜的防御反应;当若虫密度为5—6头/cm~2时,对烟蚜的抑制作用最明显,死亡率较对照升高56.8%;当烟粉虱若虫密度达11—12头/cm~2时,对烟蚜的负面影响较上一个密度略有降低。
在9—10头/cm~2烟粉虱密度条件下,随着烟粉虱若虫生长和为害时间的延长,对烟蚜防御反应的影响呈现先逐渐增强然后减弱的的现象。烟粉虱若虫为害1天时烟草白脉叶片对烟蚜没有明显影响;为害第5天,白脉叶片上烟蚜的死亡率较对照叶片升高36.1%;为害第10天烟草对烟蚜的防御反应表现最强烈,白脉叶上烟蚜的死亡率较对照叶片升高72.8%,平均相对生长率最低;为害第15天时烟粉虱若虫处于4龄拟蛹期,对烟蚜的影响较为害10天时减小,死亡率较对照植株升高58.9%。
(5)证明了烟粉虱取食诱导的烟草对烟蚜的防御反应具有明显的时空效应。采用小叶笼法研究了烟粉虱为害烟草对烟蚜死亡率、相对生长率和繁殖力的影响。研究结果表明,烟粉虱为害后的烟草植株四个叶位中,其中尤以白脉叶影响显著,影响顺序为白脉叶＞中间叶＞心叶＞虫体叶,证明了烟粉虱诱导的烟草对烟蚜的防御反应在烟草植株体内具有空间传递性。在烟粉虱发育至3龄时去除烟粉虱若虫虫体,然后接烟蚜,在去虫后15天内对烟蚜死亡率及相对生长率的影响与不去虫的对照相比均有明显差异,但随着去虫时间的延长,烟草对烟蚜的防御反应逐渐减弱;至去虫后第20天这种负面影响消失,说明烟粉虱取食诱导的烟草抗虫性能持续表达15天左右。
(6)测定了烟粉虱为害的烟草体内主要营养指标的变化。通过可溶性糖、可溶性蛋白含量变化与对烟蚜抑制影响的时间和空间分析,表明可溶性糖、可溶性蛋白的含量与烟蚜的死亡率呈密切的负相关性;受烟粉虱为害后的烟草白脉叶中氨基酸总含量上升,其中,天门冬氨酸含量降低,脯氨酸、丙氨酸、甘氨酸、缬氨酸、苯丙氨酸及赖氨酸等大多数种类含量上升。烟粉虱为害后的烟草体内总蛋白含量的变化亦与烟草抗蚜性有一定的相关。
Bemisia tabaci(Gennadius) B biotype spread rapidly when it invades in China.It has been a key pest of various vegetables,cotton,tobacco and so on.At the present time,B.tabaci B biotype occurs in tobacco farmlands of Shandong province widely.We find that there has a few Myzus persicae(Sulzer) on tobacco infested by B.tabaci when investigating in farm.It indicates that B.tabaci was predomination in competing with M.persicae.However,there have no report about the influence of defense responses induced by B.tabaci on other sap-feeding insects at home and abroad currently.We start at farming phenomenon,choose tobacco as test materials,choose B.tabaci--tobacco-M.persicae as research system,study the effect of tobacco's defense response induced by B.tabaci on population growth,development and reproduction of M.persicae,major effect factors.We also investigate induced resistance mechanism initially.Investigating defense responses of tobacco to M.persicae induced by B.tabaci have important academic meaning for definiting competitive replacement ability of B.tabaci to M.persicae,showing competitive replacement mechanism of B.tabaci to M.persicae,and implementing sustainable control effectively.Major results as follows:
(1) The effect of four tobacco varieties on development and reproduction of B.tabaci was studied.The results showed that K326,CF-965 were infested by B.tabaci easier.The developmental periods of B.tabaci were shorter than other tobacco varieties'.Survivorship and the number of eggs deposited by B.tabaci were higher than other tobacco varieties'. NC-82,NC-89 were more resistant to B.tabaci.The developmental periods of B.tabaci feeding on them were longer,survivorship were lower,and the fecundities were lower.They went against development and reproduction of B.tabaci.
(2) The effect of B.tabaci on host plant preference of M.persicae was studied with two different methods.The results showed that M.persicae adults liked to choose the plant without B.tabaci infested,compared to host plant infested by B.tabaci.And for leaves depart from host plant,tobacco leaves of different position had different effect on host plant preference of M.persicae.Compared to respective control leaves,damaged leaves had significant repellence on M.persicae,but,white-vein leaves and unexpanded leaves had significant attraction to M.persicae.
(3) Tobacco infested by B.tabaci had significant inhibition on population growth, development and reproduction of M.persicae.Mix developmental stages and nymphs of B.tabaci infested tobacco had significant influence on mortality of first generation of M.persicae.They were higher 23.8%,30.0%than that of M.persicae fed on the plant without B.tabaci infested.Compared to control plants,the population growth rate of second generation,third generation reduced 55.5%,49.2%;87.9%、79.6%,respectively.The population growth of M.persicae was inhibited significantly.The effect of B.tabaci damaged leaves and white-vein leaves on development and reproduction of M.persicae was studied with clip-cage.The results showed that the white-vein leaves infested by B.tabaci had significant detrimental influence.Compared to leaves of same position on control plant,the mortality increased 46.6%when switching on M.persicae for 8 days,developmental times prolonged 0.9 days,mean relative growth rate reduced 6.5%,and average fecundity decreased 5.7 per female.The development and reproduction of M.persicae was inhibited significantly. But,the damaged leaves had no significant negative influence on development and reproduction of M.persicae.
(4) Major effect factors of tobacco defense responses to M.persicae induced by B.tabaci were studied.The results indicated that B.tabaci nymphs were major factor which induced defenses responses to M.persicae.Defense responses of tobacco to M.persicae could not induced by B.tabaci infested 9 days with 500 adults per plant.
The effect of infested density of B.tabaci(0-1/cm~2,2-3/cm~2,5-6/cm~2,11-12/cm~2) on M.persicae was studied.The results showed that tobacco infested by B.tabaci nymphs with 0-1/cm~2 had no harmful effect on M.persicae.Tobacco defenses to M.persicae began to expression when density reached to 2-3/cm~2.When density reached to 5-6/cm~2,it was expressed the most significantly.The mortality was the highest,56.8%higher than control. When density reached to 11-12/cm~2,the negative effect on M.persicae was lower than prior infested density.
The influence of infested time of B.tabaci nymphs with 9-10/cm~2(1d,5d,10d,15d) on M.persicae was studied.Along with growing of B.tabaci nymphs and prolonging of infested time,the influence of defense responses to M.persicae began to strengthen and then weak. B.tabaci nymphs infested 1 day had no significant effect on M.persicae.Induced defense responses of tobacco began to express when nymphs infested 5 days,the mortality of M.persicae feeding on white-vein leaves was 36.1%higher than control plant.10 days infested by B.tabaci nymphs resulted in strong expression of defenses to M.persicae.The mortality of M.persicae fed on white-vein was the highest,72.8%higher than control plants. Mean relative growth rate was the lowest.B.tabaci nymphs reached to subnymph periods when they infested 15 days.The effect on M.persicae began to reduce compared to that of 10 days infested by B.tabaci nymphs,the mortality was 58.9%higher than contrast plant.
(5) Tobacco's defense response induced by B.tabaci to M.persicae had temporal and spatial effect.We studied the effect of defense responses on mortality,population growth rate, and fecundity of M.persicae with clip-cage.The results showed that among leaves of four position infested by B.tabaci,white-vein leaves had significant influence on development of M.persicae.The sequence of effect on mortality,development and reproduction of M.persicae was white-vein leaves>middle leaves>unexpanded leaves>damaged leaves.It proved that defense responses of tobacco to M.persicae can transmit spatially within tobacco.
We wiped off nymphs when B.tabaci nymphs developed to 3_(rd) instar and then switched on M.persicae.There had significant influence on M.persicae after wiped off B.tabaci nymphs for 15 days.Mortality and population growth rate of M.persicae had significant difference between treated and control plants.Along with duration prolonged,the influence began to retard.After 20 days,the negative influence disappeared.It suggested that the duration of induced resistance was around 15 days.
(6) We determinated major nutritional index in tobacco damaged by B.tabaci.The results indicated that there had negative correlation between soluble sugar,soluble protein content and mortality of M.persicae.Total content of amino acids in tobacco infested by B.tabaci increased.Content of Asp deduced,content of Pro,Ala,Gly,Val,Phe,Lys and so on increased. The total protein content of tobacco damaged by B.tabaci also had some relativity with tobacco's resistance to M.persicae.
引文
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