环境胁迫条件下麦长管蚜种下体色型分化生态机理研究
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摘要
麦长管蚜Sitobion avenae (Fabricius),属半翅目Hemiptera,蚜科Aphididae,是国内外重要的农业害虫。当今气候环境的变化对生物特别是R-生态对策的蚜虫产生着强大选择压,使其进化更为迅速。麦长管蚜有红、绿两种体色生物型,其红色型可能就是紫外线增强等诱因产生的新的种下型。种下体色生物型分化是导致麦长管蚜广泛成灾的内在因素之一。本文在前人的研究基础上,用紫外线UV-B、脱共生菌及寄主植物感染病毒(谷类黄矮病CYDV-PRV和小麦矮缩病WDV)等环境胁迫,对两种体色型麦长管蚜的生长发育、繁殖、种内竞争等生态学参数进行了研究,并利用刺探电位图谱技术(EPG)对上述环境胁迫条件下两种体色型麦长管蚜的取食行为学机制进行了研究,以期证明麦长管蚜种下体色型分化的生态机理。得到如下结果:
     1.紫外线直接照射是麦长管蚜种下体色型分化的动力之一
     在不同剂量(0,216,432,864kJ/m~2)紫外照射两种体色型麦长管蚜初生若蚜后,对其生长发育、繁殖参数和取食行为进行了测定。生长发育和繁殖参数的结果表明:低剂量紫外辐射(216kJ/m~2)能够促进两种体色型麦长管蚜的生长发育和繁殖;而高剂量紫外辐射(432,864kJ/m~2)能够抑制两种体色型麦长管蚜的生长发育和繁殖,且红色型对高剂量紫外直接照射的适应能力较绿色型强。取食行为EPG研究表明其机理在于:高剂量紫外辐射不仅抑制两种体色型麦长管蚜口针在寄主植物韧皮部的取食,还增加蚜虫口针穿过表皮/叶肉细胞的时间,且绿色型麦长管蚜口针穿过寄主植物表皮/叶肉细胞和取食韧皮部汁液的难度均高于红色型。说明红色型麦长管蚜是适应紫外增强的一种进化生物型。
     2.紫外线照射寄主植物对蚜虫的间接影响是麦长管蚜种下体色型分化的又一动力
     不同剂量(对照剂量0kJ/m~2,环境正常辐射剂量600kJ/m~2,加倍辐射剂量1200kJ/m~2)紫外照射寄主植物大麦后,对两种体色型麦长管蚜的生长发育、繁殖参数和取食行为进行了测定。生长发育和繁殖参数的结果表明:当寄主植物受到加倍强度紫外照射后,两种体色型麦长管蚜的生长发育和繁殖均受到抑制,且红色型麦长管蚜对加倍紫外强度照射后植物的适应能力较绿色型强。EPG研究结果表明其机理在于:加倍强度紫外照射寄主植物后,均能增加寄主植物对两种体色型麦长管蚜的表面抗性、表皮/叶肉抗性和韧皮部抗性,且寄主植物对绿色型麦长管蚜上述三个方面的抗性均比高于红色型。进一步说明红色型是适应紫外增强的一种进化生物型。
     3.麦长管蚜体内共生菌在其种下体色型分化过程中起着重要作用
     利用广谱性抗生素利福平脱除两种体色型麦长管蚜胞内共生菌后,对两种体色型麦长管蚜的生长发育、繁殖参数和取食行为进行了测定。生长发育和繁殖参数的结果表明:脱共生对两种体色型麦长管蚜的生长发育和繁殖都造成了不利的影响,且脱共生对红色型麦长管蚜的不利影响要比绿色型大。EPG研究结果表明其机理在于:脱共生蚜虫口针穿过植物表皮/叶肉细胞难度增加,在韧皮部的取食时间缩短,且红色型脱共生麦长管蚜上述阶段的取食难度均高于绿色型。表明蚜虫体内共生菌参与控制麦长管蚜种下体色型分化进程。
     4.寄主植物体内植物病毒对麦长管蚜的种下体色型分化有重要影响
     寄主植物大麦感染CYDV-PRV和WDV病毒后,对两种体色型麦长管蚜(非介体昆虫)的生长发育和繁殖起了促进作用,且对绿色型麦长管蚜的生长发育的促进作用较红色型强。EPG研究结果表明其机理在于:非传播性植物病毒显著减弱了植物表面、表皮/叶肉组织、韧皮部组织对两种体色型麦长管蚜的抗性,且感染植物病毒的寄主植物对绿色型韧皮部取食阶段的抗性显著低于红色型。证明寄主植物体内的植物病毒对麦长管蚜的种下体色分化有重要影响。
     5.环境胁迫影响两种体色型麦长管蚜的竞争力从而影响麦长管蚜的种下体色分化
     在环境胁迫(正常对照、加倍剂量紫外线照射、CYDV-RPV病毒)寄主植物条件下,单独和混合饲养(一头绿色型和一头红色型罩在同一微笼罩内)不同体色型麦长管蚜,分析竞争结果发现:正常饲养条件下,两种体色型麦长管蚜竞争力相当;紫外胁迫下红色型麦长管蚜的竞争力比绿色型强;而植物病毒胁迫下绿色型麦长管蚜的竞争力比红色型强。证明环境胁迫通过影响两种体色型麦长管蚜的竞争力从而影响麦长管蚜的种下体色分化,且环境胁迫因子不同其效应不同。
     总之,环境胁迫条件下两种体色型麦长管蚜适应性和竞争力具有显著差异,且胁迫因子不同其效应和生态学机理均不相同,说明环境胁迫是导致麦长管蚜种下体色型分化的主要原因。这对阐明麦长管蚜虫种下分化的机制具有重要意义,由此也可折射出R对策昆虫进化加剧的原因,并为全球环境变化加剧这一大背景下麦长管蚜和植物病毒的综合治理提供理论依据。
     本研究有以下四个方面的创新:
     1.证明紫外线、植物病毒胁迫是麦长管蚜种下体色型分化的重要动力;
     2.证明麦长管蚜体内共生菌在其种下体色型分化过程中起着重要作用;
     3.揭示出环境胁迫条件下麦长管蚜种下体色型分化的生态机理;
     4.证明紫外胁迫下红色型麦长管蚜的竞争力比绿色型强,而植物病毒胁迫下绿色型麦长管蚜的竞争力比红色型强。
Sitobion avenae (Fabricius), Hemiptera, Aphididae, is one of the most importantagricultural pests in the whole world. One of internal factors of widely harm is polarization ofbody-color morphs (red and green morphs) in species caused by the change of environment.As the selective stress, the change of environment accelerates the evolution of S. avenae.Based on previous researches, to explore the ecological mechanism of polarization ofbody-color morphs, the development, reproduction and intraspecific competition of twomorphs of S. avenae were estimated under the condition of UV-B, aposymbiosis, plant viruses(CYDV-PRV and WDV) in this study. Under these conditions, the research on feedingbehavior was also carried out by Electronical Penetration Graph (EPG). The following are theresults of this research:
     1. UV-B radiation directly is one of the driving forces for the polarization of body-colormorphs of S. avenae
     The parameters of development, fecundity and feeding behavior of S. avenae wereestimated after radiating the first instar nymphs of body-color morphs by UV-B in differentdose (0,216,432,864kJ/m~2). The results indicated that, the development and fecundity werepromoted by low-dose UV-B (216kJ/m~2), and effected negatively by high-dose UV-B (432,864kJ/m~2). For red aphids, the capacity of adaptation to this was stronger than green aphids.Trough determining the feeding behavior by EPG, we could demonstrate that: radiation ofhigh-dose UV-B not only restrained the feeding in phloem but also prolonged the prickingtime in the part of cuticle and mesophyll. The fact that the feeding behavior of green aphidspricking host plant cuticle and phloem was more difficult than that of red aphids demonstratedthat red aphids was the evolutionary biotype to adapt the UV-B.
     2. The indirect effects on S. avenae through radiating host plant by UV-B is anotherdriving force for the polarization of body-color morphs
     The parameters of development, fecundity and feeding behavior of S. avenae wereestimated after radiating barley by UV-B in different dose (0kJ/m~2as the control,600kJ/m~2as normal UV-B radiation, double-dose UV-B is1200kJ/m~2). The results indicated that: thedevelopment and fecundity of two body-color morphs were restrained significantly under thecondition of radiating post plant by double-dose UV-B. It could include that two types of aphids had adapted post plants exposed to the normal UV-B radiation. For Red aphids, thecapacity of adaptation to this was stronger than green aphids. The result of feeding behaviorcould indicate that cuticle resistance, mesophyll resistance and phloem resistance of two typesof S. avenae were all increased under the condition of double-dose UV-B radiation. Moreover,the resistance of post plant to green aphids was higher than the Red aphids exposed to threetreatments, which further demonstrated that red aphids demonstrated that red aphids was theevolutionary biotype to adapt the UV-B.
     3. Aposymbiosis in S. avenae plays an important role in the polarization course ofbody-color morphs
     The parameters of development, fecundity and feeding behavior of S. avenae wereestimated after disrupting symbiotic bacteria from aphids by rifampicin which was one kindof antibiotics. The results indicated that: the development and fecundity were influencednegatively for the two morphs Bacteria-free aposymbiotic aphids. The influence ofaposymbiosis to Red aphids was more serious than green aphids. It was demonstrated fromthe result of feeding behavior that the feeding of two morphs aphids was restrained, such as,pricking time in the part of cuticle and mesophyll was prolonged and the time of feeding inphloem was shortened. Furthermore, the feeding behavior of Red aphids was more difficultthan that of green aphids. It was indicated that aposymbiotic involved in the polarizationcourse of body-color morphs of S. avenae.
     4. The non-transmission plant virus could affect polarization of body-color morphs of S.avenae
     The development and fecundity of two body-color aphids could be promoted afterinfecting host plant (barley) successfully by non-propagative CYDV-PRV and WDV. Theeffect on green aphids was stronger than the red aphids. The result of feeding behavior couldindicate that cuticle, mesophyll and phloem resistance to two types of S. avenae were alldecreased by non-propagative CYDV-PRV and WDV. Moreover, the phloem resistance ofhost plant infected viruses to green aphids was lower than the red aphids. Therefore thenon-transmission plant virus plays an important role in the polarization course of body-colormorphs.
     5. The population competitiveness of two body-color morphs of S. avenae induced byenvironmental stress could influence polarization of body-color morphs
     The parameters of development and fecundity of S. avenae were estimated through risingseparately and mixed in the host plants exposed to the treatments of control, double-doseUV-B radiation and CYDV-RPV virus. The results indicated that: in the treatment of control,the competitiveness of two body-color morphs had not significant difference. UV-B radiation could generate stronger competitiveness for green aphids. However, the red aphids had highercompetitiveness under the induction of plant viruses. It could demonstrate that environmentalstress could influence polarization of body-color morphs through affecting populationcompetitiveness of two body-color morphs of S. avenae.
     In short, the adaptation and competitiveness of two body-color morphs of S. avenae weredifferent significantly under the environmental stress. The effect and the ecologicalmechanism were also various based on the different environmental factors. We candemonstrate that environmental stress is the main cause of polarization of body-color morphs,which has the important significance for exploring the mechanism of polarization and providethe theoretical basis for the management of S. avenae and plant virus under the condition ofever-changing environment.
     The innovations are following in this study:
     1. We demonstrate that UV-B radiation and plant virus were the driving forces for thepolarization of body-color morphs of S. avenae.
     2. It was demonstrated that aposymbiosis in S. avenae played an important role in thepolarization course of body-color morphs.
     3. The ecological mechanism of polarization of body-color morphs of S. avenae wasproposed.
     4. The conclusion that UV-B radiation could generate stronger competitiveness for greenaphids and the red aphids had higher competitiveness under the induction of plant viruses wasobtained in this research.
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