日本刀角瓢虫对不同叶毛密度茄子品种的产卵选择性

【目的】产卵是昆虫发育过程中繁衍后代、维持种群数量的重要环节。在前期研究发现日本刀角瓢虫Serangium japonicum喜欢在茄子Solanum melongena叶片上产卵的前提下,本研究旨在深入探究日本刀角瓢虫对不同叶毛密度茄子品种的产卵选择性,以期进一步明确植物叶毛介导的日本刀角瓢虫产卵选择机制。【方法】观察日本刀角瓢虫雌成虫在4个茄子品种(黑到底、黑妞、日友长直壮、络星)叶碟和植株上的产卵选择性,测定不同茄子品种上日本刀角瓢虫后代的生长发育及成虫的繁殖力、捕食能力和对叶片的附着力,观察4个茄子品种叶片背面超微结构,测定日本刀角瓢虫雌成虫对茄子叶片的气味选择,测量茄子叶片上烟粉虱Bemisia tabaci卵粒的长度和宽度,测定日本刀角瓢虫卵被其雌成虫的同类捕食风险。【结果】日本刀角瓢虫偏好在黑到底上产卵,后代个体在黑到底品种上总发育历期(15.41±0.21 d)最短,亲代单雌产卵量(877.44±15.27粒/头)最高。但是,日本刀角瓢虫的产卵量占比与子代生长表现(发育历期、存活率和成虫体重)和成虫生存表现(繁殖力和捕食能力)均无显著相关性,而与叶片背面叶毛密度和长度存在显著正相关性;日本刀角瓢虫在茄子叶碟上的产卵量占比与日本刀角瓢虫雌成虫附着力存在显著正相关性。日本刀角瓢虫雌成虫附着力与叶毛密度显著正相关,而与亲代生长表现无显著相关关系。茄子叶片气味、猎物质量以及被同类捕食风险均不显著影响日本刀角瓢虫的产卵选择。【结论】茄子叶片背面叶毛密度及叶毛介导的日本刀角瓢虫附着力在日本刀角瓢虫对茄子叶片的产卵选择性中起重要作用。     

冷藏温度对日本刀角瓢虫存活及生长发育的影响

【目的】本研究旨在调查不同冷藏温度下日本刀角瓢虫Serangium japanicum成虫生物学特性及F₁代的生长发育,明确日本刀角瓢虫成虫最适冷藏温度。【方法】将日本刀角瓢虫成虫置于不同低温(7,10,13和16℃),贮藏10 d时测定其存活率、单雌产卵量、寿命和对烟粉虱Bemisia tabaci 4龄若虫的日捕食量,以及日本刀角瓢虫F₁代存活率和发育历期;qRT-PCR测定日本刀角瓢虫成虫体内热激蛋白基因Hsp70和Hsp90的相对表达量。【结果】日本刀角瓢虫成虫在16℃低温贮藏10 d时,其存活率、雌雄成虫寿命、单雌产卵量和日捕食量以及F₁代存活率均与贮藏在26℃的对照无显著差异(存活率：99%vs 100%;雌成虫寿命：110.65 d vs 106.87 d;雄成虫寿命：123.12 d vs 108.79 d;单雌产卵量：399.19粒vs 422.63粒;日捕食量：34.70头vs 31.95头;F₁代存活率：80.39%vs 94.12%);但其F₁代发育历期...     

烟粉虱天敌日本刀角瓢虫的捕食行为

研究了日本刀角瓢虫雌成虫和幼虫捕食粉虱时的各种行为时间分配.结果表明,日本刀角瓢虫成虫的捕食行为可分为爬行、取食、清洁、静息、整翅和排泄6个部分.饥饿后的日本刀角瓢虫幼虫捕食烟粉虱卵时,幼虫各龄期间的取食和爬行时间无显著差异,对烟粉虱卵的处置时间随瓢虫幼虫虫龄的增大而缩短.日本刀角瓢虫雌成虫捕食烟粉虱卵时,饥饿后的雌成虫用于取食的时间显著长于非饥饿的雌成虫,而用于清洁、静息和爬行的时间显著短于非饥饿的雌成虫;饥饿后的瓢虫雌成虫对卵的处置时间显著长于非饥饿的雌成虫.饥饿后的日本刀角瓢虫雌成虫捕食烟粉虱若虫时,瓢虫雌成虫的取食时间随烟粉虱若虫虫龄的增大而增加,清洁和静息时间随烟粉虱若虫虫龄的增大而缩短,对若虫的处置时间随烟粉虱若虫虫龄的增大而增加.     

日本刀角瓢虫形态特征及生物学特性研究

日本刀角瓢虫是粉虱类害虫的重要捕食性天敌之一.本文详细地描述了日本刀角瓢虫的形态特征,并附有各虫态的特征图.且对日本刀角瓢虫的生活习性进行了观察,以烟粉虱卵为食料,日本刀角瓢虫世代发育历期平均为22.56天,成虫平均寿命为91.94天,平均每雌产卵量为564.8粒;以烟粉虱若虫为食料,日本刀角虫世代发育历期平均为21.70天,成虫平均寿命为81.62天,平均每雌产卵量为650.0粒;瓢虫可捕食烟粉虱的卵、若虫、"蛹"、成虫各虫态,是烟粉虱的有效天敌,具有很好的保护利用价值.     

粉虱小毛瓢虫Nephaspis oculatus(鞘翅目,瓢虫科)捕食功能反应与寄主植物选择(英文)

Nephaspisoculatus (Blatchley)属毛瓢虫亚科的小毛瓢虫族 ,专食粉虱。它源于中美洲 ,自七十年代初就在美国佛州被发现 ,是柑橘和蔬菜上白粉虱的重要天敌。本研究确定了该瓢虫各幼虫龄期和雌雄成虫在青菜 (col lard)叶上捕食银叶白粉虱BemisiaargentifoliiBellows&Perring卵在 6个密度的捕食功能反应及其成虫对 5种寄主植物的选择。结果表明 ,该瓢虫捕食率与白粉虱卵的密度呈线性相关。除第四龄幼虫其相关系数 (r2 )仅为0 .6 33外 ,其它各虫态的相关系数高达 0 .82 7～ 0 .94 4。根据该瓢虫各龄幼虫和雌雄成虫捕食量和白粉虱卵密度的关系 ,其捕食功能反应符合Holling圆盘方程的第二种反应模式。随着龄期的增加 ,幼虫捕食白粉虱卵的量也随之增加。第四龄幼虫捕食白粉虱卵量最多可达 312粒 ,其次为第三龄幼虫 2 2 9粒 ,第二龄幼虫 73粒 ,而第一龄幼虫捕食量仅为 34粒。雌成虫捕食白粉虱卵量 (32 1粒 )明显高于雄成虫 (2 37粒 )。雌成虫特别喜欢在青菜上活动、取食和产卵 ,而不喜欢番茄 ,大豆 ,茄子和甘薯。有 4 4 .9%的雌成虫在青菜上活动 ,只有 9.9% - 17.1%的雌成虫在其它植物上活动。雌成虫在青菜 ,甘薯和茄子上的产卵量占总产卵量的比率依次为 95 .8% ,2 .8%和 1.4 % ,而无任何卵产在番茄和大豆上。     

烟粉虱成虫对黄瓜叶龄的喜好性及与后代发育适合度间的关系

在室内采用笼罩和发育测定方法,研究了烟粉虱成虫对黄瓜叶龄的喜好性及其与后代生长发育、存活之间的关系,同时调查了叶毛密度与成虫喜好性之间的相关关系。结果表明(1)烟粉虱成虫比较喜欢在黄瓜植株的中部叶片取食和产卵。(2)叶龄对烟粉虱的存活率没有显著影响,对烟粉虱的发育历期只在卵期和若虫四龄期表现出显著影响,对其他龄期或整个世代均没有显著影响。(3)叶毛密度随着叶龄的增大而顺次显著减小;在接虫初始阶段,烟粉虱成虫的叶片选择性与叶毛密度成显著负相关关系,但72h后具有中等叶毛密度的叶片上的成虫数量和着卵量均显著高于叶毛密度较大或较小的叶片。研究结果对培育抗烟粉虱的黄瓜品种和改进烟粉虱的防治措施具有一定指导意义。     

红点唇瓢虫对矢尖蚧的捕食作用(英文)

红点唇瓢虫（Chilocoruskuwanae）的产卵雌虫对矢尖蚧（Unaspisyanonensis）的捕食量显著大于雄性成虫。在25℃下,雌成虫日平均捕食矢尖蚧雌成蚧42．7头,而雄成虫的日捕食量仅为22．3头。但48h饥饿处理可消除此差异。要使雌成虫产卵,每天必须至少捕食15头矢尖蚧的雌成蚧。红点唇瓢虫成虫对矢尖蚧各虫态的功能反应均为HollingⅡ型。根据对雌成蚧的功能反应,在16－35℃范围内最大捕食量随温度的升高而升高,在37℃下则急剧下降。温度并不改变功能反应的类型。由红点唇＊虫成虫对雌成钛的捕食作用,估算得最低临界攻击温度、最佳攻击温度和最高临界攻击温度分别为10．58℃、31．53℃和3820℃。捕食空间增大或捕食者数量的增加都会使攻击率下降;但是捕食者密度变大时,捕食空间的影响将大大变小。红点唇瓢虫成虫对矢尖蚧的雄蛹表现出最大的猎物选择性。它们对矢尖蚧各虫态的捕食选择性依次为：雄蛹、2龄雄蚧、2龄雌蚧和1龄若虫。     

瓜蚜寄主植物对多异瓢虫捕食功能反应的影响

本研究比较了瓜蚜Aphis gossypii Glover 5种主要寄主植物对多异瓢虫Hippodamia variegta(Goeze)捕食功能反应的影响。结果表明,瓜蚜寄主植物对多异瓢虫功能反应有显著的影响。在大多数猎物密度下, 1~3龄幼虫对南瓜和黄瓜上瓜蚜的捕食量较瓢葫芦和哈密瓜上的高;在蚜虫密度较低时（≤ 50头/皿）,4龄幼虫和成虫对南瓜和黄瓜上瓜蚜的捕食量较高,而蚜虫密度≥ 70头/皿时,对瓢葫芦和搅瓜上瓜蚜的捕食量较高。多异瓢虫各龄幼虫和成虫对5种寄主植物上瓜蚜的功能反应均符合Holling II反应。随着瓢虫龄期的增大,猎物处理时间缩短,瞬时攻击率增大,最大捕食量增加。多异瓢虫成虫处理猎物的时间较长于4龄幼虫,而瞬时攻击率和最大捕食量小于4龄幼虫。多异瓢虫捕食不同寄主植物上饲养的瓜蚜时,猎物处理时间、瞬时攻击率和最大捕食量不同。     

七星瓢虫卵的低温贮藏条件研究

为了探究七星瓢虫卵的最适低温贮藏条件,本试验研究了七星瓢虫卵在5、7、9和11℃条件下,分别贮藏10、 20和30 d的孵化情况,结果表明七星瓢虫卵在11℃贮藏10 d条件下孵化率最高,可以达到50.67%,进一步分析最适的低温贮藏条件11℃贮藏10 d对七星瓢虫种群适合度的影响。结果显示11℃贮藏10 d对七星瓢虫化蛹率无显著影响。以豌豆修尾蚜为猎物,对其幼虫和成虫的日捕食量进行检测,结果表明七星瓢虫的卵经11℃贮藏10 d后,各个龄期幼虫和成虫的日捕食量与对照组相比无显著差异。但经11℃贮藏10 d后七星瓢虫的发育历期显著缩短,羽化率、产卵量以及子代孵化率显著降低。上述研究结果对利用低温环境提高七星瓢虫的贮藏效率提供一定的理论基础。     

异色瓢虫非同胞间食卵自相残杀对成虫生殖力的影响

【目的】食卵自相残杀现象常见于瓢虫中,业已明确取食同胞卵对于初孵瓢虫至关重要,但取食非同胞卵的营养价值是否高于蚜虫仍存争议。本研究旨在明确异色瓢虫Harmonia axyridis在幼虫期除取食蚜虫外兼捕食非同胞卵是否影响其雌成虫的生殖表现。【方法】在异色瓢虫幼虫期设置3个饲喂处理,即仅饲喂禾谷缢管蚜Rhopalosiphum padi(对照)、除饲喂蚜虫外还分别给各龄幼虫饲喂少量非同胞卵(分散食卵)以及除饲喂蚜虫外仅给高龄幼虫饲喂较多非同胞卵(集中食卵)等,观察雌成虫体型大小、产卵前期、连续10 d的逐日产卵量和卵孵化率等形态和生殖特性。【结果】饲喂处理对瓢虫雌成虫体型大小无显著影响,对照、分散食卵和集中食卵3种饲喂处理下的雌成虫身体体积分别为92.30,89.73和93.98 mm~3;饲喂处理对雌成虫产卵前期也无显著影响,3种饲喂处理下雌成虫产卵前期分别为7.50,7.74和7.70 d;但饲喂处理对雌虫生殖力具有显著影响,分散食卵处理与对照无显著差异,羽化后10 d分散食卵和对照下累计产卵量分别为185和205粒,但集中食卵(88粒)比对照或分散食卵的产卵量显著降低;饲喂处理对卵孵化率无显著影响,对照、分散食卵和集中食卵3种饲喂处理下的卵孵化率分别为63.10%,72.58%和66.61%。【结论】结果说明,异色瓢虫各龄幼虫偶尔取食非同胞卵不影响其成虫的生长发育和生殖,但高龄幼虫捕食较多非同胞卵可降低其雌成虫的生殖力。     

Influence of trichomes on attractiveness and ovipositional preference of Bemisia tabaci (Genn.) B biotype (Hemiptera: Aleyrodidae) on tomato genotypes

Brazil is one of the world's largest tomato producer, but considerable part of the production is lost due to the attack of Bemisia tabaci (Genn.) B biotype. Resistant germoplasm plants can be an important method for B. tabaci control in integrated pest management approaches. The attractiveness and ovipositional preference of B. tabaci for 17 tomato genotypes were evaluated in a free-choice test. Trials were set up in a randomized block design with ten replicates. Each replicate (one tomato plant per pot) was placed in a cage (80 x 50 x 50 cm) and infested with 1,000 adults during four days. Linear correlation tests were applied between the number of insects and eggs and number of trichomes in each tomato genotype. LA716, LA444-1 and PI134418 genotypes were the least attractive, while the 'Santa Clara' was the most attractive; PI134417 trapped the largest number of adults. LA716 genotype (4.1 eggs/leaflet and 2.1 cm2/eggs per leaflet) was the least preferred for whitefly oviposition; NAV1062, 'Fanny', LA1335, 'Santa Clara' and IAC294 were the most preferred genotypes. The glandular trichomes density was negatively correlated with whitefly's attractiveness and oviposition per leaflet and per leaf, and positively with the number of trapped insects. The non-glandular trichomes density was negatively correlated with the number of trapped insects and positively with whitefly's oviposition per cm2/leaflet andper cm2/leaf. LA716 had high antixenosis level (ovipositional nonpreference) toward B. tabaci B biotype related with type IV glandular trichome.     

Fecundity and egg distribution of the herbivore Spodoptera pectinicornis as influenced by quality of the floating aquatic plant Pistia stratiotes

We determined the influence of larval host quality of the floating aquatic weed Pistia stratiotes L. (Araceae) on the fecundity and egg distribution of the biological control agent Spodoptera pectinicornis (Hampson) (Lepidoptera: Noctuidae). Different nutritional levels were produced by growing plants with relatively low and high levels of nitrogen, phosphorus and potassium fertilizer. Female and male pupal biomass were significantly greater when the larvae were fed leaves from the high fertilizer plants. Although the fertilizer treatments did not significantly influence total fecundity, there was an indirect effect as adults from larger pupae were more fecund. Regardless of treatment, ovipositing females formed a depression in the leaf surface by removing trichomes into which they deposited the egg masses. Most of the egg masses were laid on the lower leaf surface, on leaf positions 5–8 (counting from the young inner to the outer leaves) and during days 1–2 post-eclosion. Most of the eggs were laid in masses but about 12% were solitary. Females fed the low fertilizer treatment laid a greater proportion (mean ± s.e.) of their eggs as solitary eggs (17.3 ± 3.4% of total eggs) than did females fed the high fertilizer treatment (8.3 ± 2.3% of total eggs). The increased percentage of solitary eggs laid by the females from the low quality larval diet may be an adaptive response to decrease competition among the progeny.     

Influence of Canopy Height and Fertilization Levels on the Resistance of Lycopersicon Hirsutum to Aculops Lycopersici (Acari: Eriophyidae)

The objective of this work was to study the effect of NK fertilization levels and canopy height on the resistance of Lycopersicon hirsutum and Lycopersicon esculentum to Aculops lycopersici (Acari: Eriophydae). The effects of NK fertilization levels and canopy height in the leaf size and density of trichomes and their effects on tridecan-2-one (2-TD) and undecan-2-one (2-UD) limiting the attack of A. lycopersici on tomato plants were assessed. Different NK fertilization levels had no effect on the resistance of L. hirsutum to A. lycopersici. No significant differences were found in attack rates of this mite on leaves of the top and median parts of L. hirsutum canopy. The type and density of trichomes were the main determining factor of A. lycopersici attack on tomato plants. High trichome densities and type VI glandular trichomes which produce tridecan-2-one are important resistance factors on tomato plants. L. hirsutum showed a high resistance level to A. lycopersici due to high densities of type VI glandular trichomes and consequently higher levels of tridecan-2-one in its leaves.     

Inhibition of Telenomus sphingis an egg parasitoid of Manduca spp. by trichome/2-tridecanone-based host plant resistance in tomato

The resistance of accession PI 134417 of the wild tomato Lycopersicon hirsutum f. glabratum C. H. Mull to Manduca sexta (L.) (Lepidoptera: Spingidae) and Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae) is conditioned by the high densities of 2-tridecanone-containing, glandular trichomes associated with the foliage. In laboratory experiments, rates of parasitism of M. sexta eggs by Telenomus sphingis (Ashmead) (Hymenoptera: Scelionidae) were lower among eggs on PI 134417 foliage than among eggs on foliage of the cultivated tomato L. esculentum Mill. (cv. Better Boy). The latter is characterized by a significantly lower density of type VI glandular trichomes than PI 134417 and an absence of 2-tridecanone. Parasitism by T. sphingis was also reduced among eggs on foliage of the F₁ hybrid between PI 134417 and L. esculentum. The hybrid foliage lacks 2-tridecanone but has a density of type VI glandular trichomes that is intermediate between those of PI 134417 and L. esculentum, indicating that elevated densities of type VI glandular trichomes adversely affect T. sphingis. This conclusion was further substantiated by the finding that there were no differences among plant lines in the levels of parasitism of M. sexta eggs when the eggs were on foliage that had been divested of glandular trichomes.In bioassays in which T. sphingis adults or immatures in host eggs were exposed to filter paper treated with 2-tridecanone at rates comparable to those associated with PI 134417 foliage, 2-tridecanone was acutely toxic and caused high levels of mortality. In addition, at high concentrations, 2-tridecanone vapors were repellent to T. sphingis adults. However, when exposed to PI 134417 foliage, few T. sphingis adults were killed.Parasitism of M. sexta eggs was unaffected when the eggs were deposited by moths reared as fifth instar larvae on diet containing 2-tridecanone and/or 2-undecanone at levels comparable to those associated with PI 134417 foliage.     

ToCV单独侵染和TYLCV&ToCV复合侵染番茄植株上烟粉虱寄主适应性及寄主植物营养成分含量和防御反应变化

[目的]本研究以番茄褪绿病毒(tomato chlorosis virus,ToCV)和番茄黄化曲叶病毒(tomato yellow leaf curl virus,TYLCV)为主体,旨在明确ToCV单独侵染及TYLCV&ToCV复合侵染对烟粉虱Bemisia tabaci MED隐种寄主适应性的影响,并从寄主植物营...     

番茄潜叶蛾卵黄原蛋白受体基因TaVgR在生殖发育调控中的作用

【目的】本研究旨在明确卵黄原蛋白受体(vitellogenin receptor, VgR)在番茄潜叶蛾Tutaabsoluta生殖发育过程中的功能,为潜叶类害虫的绿色防控提供候选靶标。【方法】基于番茄潜叶蛾转录组数据,采用RT-PCR扩增TaVgR基因cDNA全序列,并进行生物信息分析;通过RT-qPCR分析TaVgR在番茄潜叶蛾不同发育阶段(1-4龄幼虫、1-7日龄雌蛹和雌成虫)和雌成虫不同组织(头、体壁、前肠、中肠、后肠、卵巢、脂肪体和马氏管)中的表达模式;进一步利用RNAi抑制番茄潜叶蛾雌蛹体内TaVgR的表达,并观测沉默TaVgR基因后番茄潜叶蛾卵巢发育及繁殖力的变化。【结果】克隆获得番茄潜叶蛾TaVgRcDNA(GenBank登录号: MZ682118)序列,其开放阅读框序列长5 496 bp,编码1 831个氨基酸,推测的蛋白分子量约为206 kD,等电点为5.17,信号肽包含N-端前18个氨基酸残基,并具有典型LDLR家族蛋白保守功能域。RT-qPCR结果显示,TaVgR转录水平随着番茄潜叶蛾龄期的增加逐渐上升,雌成虫羽化后达到最高水平;TaVgR在番茄潜叶蛾雌成虫的卵巢中表达量最高。TaVgR RNAi对初期雌蛹中TaVgR的表达抑制率为62.04%~72.55%,导致卵黄蛋白在卵巢中的沉积受阻,卵巢管和卵粒长度缩短,成虫10日单雌总产卵量及后代卵孵化率降低,最终引起番茄潜叶蛾繁殖力下降。【结论】TaVgR基因在番茄潜叶蛾雌成虫和卵巢中高表达,且沉默该基因严重阻碍其卵巢发育和降低繁殖力。本研究为开发以VgR基因作为靶标的鳞翅目害虫防治新技术奠定了理论基础。     

室内条件下以枣树为寄主的苹褐带卷蛾的生长发育、繁殖及对产卵基质的选择性

【目的】苹褐带卷蛾Adoxophyes orana在陕北红枣种植区对枣树Ziziphus jujube的危害有逐年加重的趋势。本研究旨在明确以枣树为寄主的苹褐带卷蛾发育历期、繁殖能力及雌虫对产卵基质的选择性,为准确预测、高效治理该虫提供理论依据。【方法】在室温25±1℃、相对湿度70%±5%、光周期15L∶9D的条件下,以枣树叶片为食料,观察苹褐带卷蛾实验种群的生长发育及繁殖能力;通过测定补充不同营养物质后雌蛾的单雌产卵量,评价外源营养对苹褐带卷蛾繁殖力的影响;通过观察苹褐带卷蛾雌蛾在硫酸纸、滤纸、白纸、PE保鲜膜及装订胶片5种基质上的总落卵量,确定其对无机产卵基质的选择性。【结果】苹褐带卷蛾卵、幼虫、蛹和成虫的平均发育历期分别为7.38±1.22, 16.59±2.16, 7.01±0.79和16.65±5.15 d,平均世代历期为33.87±3.64 d。苹褐带卷蛾在枣树上的内禀增长率(rm)、周限增长率(λ)和种群加倍时间(Dt)分别为0.15 d^-1,1.16 d^-1和4.67 d,平均产卵量达339.52±129.93粒/雌。雌虫可多次产卵,表现出明显的昼夜节律(多在0:00-8:00时段内);平均产卵次数6.26±2.09次,雌虫羽化后第3-5天达到产卵盛期。雌蛾在PE保鲜膜上的总落卵量显著大于在其他产卵基质上的总落卵量。补给不同浓度白糖水的雌蛾,其产卵量与补给纯净水的雌蛾的产卵量差异不显著。【结论】苹褐带卷蛾能够以枣树叶片为食物完成发育并繁殖可育后代,PE保鲜膜是其雌蛾适宜的无机产卵基质。     

Differences in distribution and performance of two sap-sucking herbivores on glandular and non-glandular Datura wrightii

1. Datura wrightii Regel (Solanaceae) is polymorphic with regard to trichome type. Some plants are densely covered with short, non-glandular trichomes, whereas other plants in the same populations possess glandular trichomes that excrete a sticky exudate. The hypothesis that glandular trichomes enhance resistance to all small insect herbivores is evaluated.
2. Field censuses in four southern Californian D. wrightii plant populations revealed that glandular plants are indeed resistant to whitefly spp. (Homoptera: Aleyrodidae). Whiteflies are almost exclusively found on non-glandular plants. In contrast, Tupiocoris notatus (Distant) (Heteroptera: Miridae), another sap-sucking herbivore of similar body size, is found predominantly on plants with glandular trichomes.
3. Laboratory experiments showed that whiteflies are unable to colonize glandular D. wrightii phenotypes. After the whitefly adults had landed on the leaves of these plants, they were trapped in the exudate and died.
4. Tupiocoris notatus adults, on the other hand, laid significantly more eggs on glandular plants. The presence of the exudate was shown to be the cue that determined their choice of glandular plants.
5. In no-choice experiments, T. notatus nymphs reared on glandular plants had significantly higher survival rates and had shorter developmental periods than those raised on non-glandular plants. This, combined with the higher oviposition rates, resulted in higher T. notatus population growth rates on glandular plants than on non-glandular plants.
6. Glandular trichomes are not therefore a universal protection against small herbivores. Differences in distribution over the two plant types within the natural herbivore guild on D. wrightii may, among other selection pressures, contribute to the maintenance of the observed trichome polymorphism.     

Comparative analysis of leaf trichome structure and composition of epicuticular flavonoids in Finnish birch species

The morphology, ultrastructure, density and distribution of trichomes on leaves of Betula pendula, B. pubescens ssp. pubescens, B. pubescens ssp. czerepanovii and B. nana were examined by means of light, scanning and transmission electron microscopy. The composition of flavonoids in ethanolic leaf surface extracts was analysed by high pressure liquid chromatography. All taxa examined contained both glandular and non-glandular trichomes (short and/or long hairs) but differed from each other in trichome ultrastructure, density and location on the leaf. Leaves of B. pubescens were more hairy than those of B. pendula, but the latter species had a higher density of glandular trichomes. Of the two subspecies of B. pubescens, leaves of ssp. pubescens had more short hairs on the leaf surface and four times the density of glandular trichomes of leaves of ssp. czerepanovii, whereas, in the latter subspecies, short hairs occurred largely on leaf veins, as in B. nana. The glandular trichomes were peltate glands, consisting of medullar and cortical cells, which differed structurally. Cortical cells possessed numerous small, poorly developed plastids and small vacuoles, whereas medullar cells had several large plastids with well-developed thylakoid systems and fewer vacuoles. In B. pubescens subspecies, vacuoles of the glandular cells contained osmiophilic deposits, which were probably phenolic, whereas in B. pendula, vacuoles of glandular trichomes were characterized by the presence of numerous myelin-like membranes. The composition of epicuticular flavonoids also differed among species. The two subspecies of B. pubescens and B. nana shared the same 12 compounds, but five of these occurred only in trace amounts in B. nana. Leaf surface extracts of B. pendula contained just six flavonoids, three of which occurred only in this species. In summary, the structure, density and distribution of leaf trichomes and the composition of epicuticular flavonoids represent good taxonomic markers for Finnish birch species.