Host tolerance does not impose selection on natural enemies

Coevolution between hosts and their natural enemies is believed to operate through the evolution of resistance traits. Although the importance of tolerance to natural enemies as an alternative defensive strategy has been recognized, there is still no consensus about the possible role of host tolerance in the evolutionary outcome of the interaction. Here, using bioassay experiments, we tested the hypothesis that variation in host tolerance among selected plant genotypes could impose a selection pressure upon a specialist herbivore. Tolerance did not affect herbivore larvae survival, weight gain, efficiency of food consumption, total food consumption, developmental time and adult mass. These results therefore do not support the hypothesis that host tolerance could affect natural enemy performance. However, resistance did negatively affect herbivore larva survival. Genetic variation in herbivore larva survival was detected, thus suggesting the potential for a coevolutionary response. Our results indicate that host tolerance would reduce opportunities for a coevolutionary response by the natural enemies of the host. Contrary to predictions from previous models, our results suggest that host tolerance may constitute an evolutionarily unstable defensive strategy.     

Divergence of Eurosta solidaginis in response to host plant variation and natural enemies

We tested the hypothesis that forest and prairie populations of the gall-inducing fly, Eurosta solidaginis, have diverged in response to variation in selection by its host plant Solidago altissima, and its natural enemies. A reciprocal cross infection design experiment demonstrated that fly populations from the prairie and forest biomes had higher survival on local biome plants compared to foreign biome host plants. Flies from each biome also had an oviposition preference for their local plants. Each fly population induced galls of the size and shape found in their local biome on host plants from both biomes indicating a genetic basis to the differences in gall morphology. Solidago altissima from the prairie and forest biomes retained significant morphological differences in the common garden indicating that they are genetically differentiated, possibly at the subspecies level. The populations are partially reproductively isolated as a result of a combination of prezygotic isolation due to host-associated assortative mating, and postzygotic isolation due to low hybrid survival. We conclude that E. solidaginis is undergoing diversifying selection to adapt to differences between prairie and forest habitats.     

Costs of resistance to natural enemies in field populations of the annual plant Arabidopsis thaliana

The annual plant Arabidopsis thaliana is widely used as a model system in molecular genetics, but little is known about populations in the field. In this experimental field study of natural populations of Arabidopsis, I tested the assumption that plant resistance has fitness costs. Models of the evolution of resistance assume a cost, which is envisioned as a reduction in fitness in the absence of natural enemies, such as insect herbivores and pathogens. The presumed basis of this cost is the diversion of limiting resources away from present and future growth and reproduction. Recent failures to detect allocation costs of resistance to herbivores have raised questions about whether costs exist and, thus, about the appropriateness of theories that postulate such costs. I found genetic variation for two traits commonly thought to function as resistance characters: trichome density and total glucosinolate concentration. Under field conditions, these characters both reduced damage by the natural assemblage of herbivores and exhibited significant fitness costs.     

Parasitoid exploitation of the seasonal variation in host plant volatile emission for herbivore location

Volatile compounds from the apple, Malus domestica Borkh. (Rosaceae), change considerably as the season progresses, and this is successfully exploited by the female codling moth Cydia pomonella L. (Lepidoptera, Tortricidae), as it searches for oviposition sites. In this study, we investigated the effect of seasonal emissions of apple fruit volatiles on the host location behaviour of a parasitoid of the codling moth larvae, Hyssopus pallidus (Askew) (Hymenoptera: Eulophidae). In dual choice olfactory bioassays, the behaviour of the parasitoid in response to apple cues was observed over the complete 2003 growing season. Our results show that codling moth infested apples evoked a strong response from the parasitoid at the beginning of the season, until July. Then, attraction dropped drastically, increasing again at the beginning of August. At the end of the growing season, just prior to harvest, infested apples hardly evoked any behavioural response. Interestingly enough, mid‐season emissions of healthy apples were per se attractive to the parasitoid, and even preferred over volatiles from infested apples. Simultaneous volatile collections from healthy apples on twigs in the field were analysed throughout the season, showing that the overall quantity of headspace volatiles peaks at the beginning of June and mid‐August. The seasonal volatile emission is correlated with the behaviour of the parasitoid during the fruit ripening stage. The results are discussed in relation to the use of H. pallidus as a potential biocontrol agent, in order to enhance current integrated pest management (IPM) programs.     

Contributions of climate,plant phenology and natural enemies to the seasonal variation of aphids on cabbage

Brevicoryne brassicae (L), Lipaphis erysimi (Kalt) and Myzus persicae (Sultzer) (Homoptera: Aphididae) form the aphid complex that causes great losses in Brassicaceae in tropical and subtropical regions. Knowledge of their population dynamics is important for the development of integrated pest management programmes. This study aimed to investigate the effects of cabbage phenology, climatic factors and natural enemies populations on the dynamics of these organisms, and the factors regulating their predators’ occurrence. The densities of aphids and their natural enemies and the climate were monitored for two years in 16 cabbage crops. The highest densities of the aphids occurred during periods of relative humidity (RH) drop, a condition that affects them positively. Regarding the predators, the factors affecting their abundance varied but RH was positively related to most of them. This study provides relevant insights into the factors that regulate the aphids in cabbage and for the decision‐making process of control of these severe pests.     

Rhythm of honeydew excretion by the tea aphid and its attraction to various natural enemies

The rhythm of honeydew excretion by the tea aphid Toxoptera aurantii (Boyer) and its attraction to following 9 species (or subspecies) of beneficial insects, Aphidius sp., Chrysopa sinica Tjeder, Chrysopa septempunctata Wesmael, Sphaerophoria menthastri L., Coccinella septempunctata L., Leis axyridis (Pallas) ab. bimaculata Hemmelmann, L. axyridis (Pallas) ab. conspicua Faldermaenn, L. axyridis (Pallas) var. spectabilis Faldermaenn and L. axyridis (Pallas) var. novemdecimpunctata Faldermaenn, were investigated. Forty-five wingless virginoparae nymphs, reproduced by the same wingless virginogenia adult within 1 h, were introduced onto tea seedlings with one aphid per seedling. The honeydews excreted from different instars of nymphs and adults were collected under 21°C, 85% RH, 3500 lx and 12 L:12 D photoperiod. It took 32.4 d ±5.8 d for the tested tea aphids to complete the development of their nymph and adult stages, during which 325.6±35.8 droplets (ca. 41.98 μl ±6.14 μl and 45.34 mg ± 8.76 mg) of honeydews were secreted. During the 1st and 4th instars, there was a logistic regression relationship between the amount of honeydews excreted and the time (days). The honeydew secretion during the first two instars was less than that from the later instars. Adult aphids survived for 22.0 d ± 0.0 d, and excreted 176.31 ± 22.38 droplets (ca. 30.38 μl ± 5.32 μl) of honeydews in a rate of 1 drop per ca. 30–50 min for 5–8 h with a pause for 2–5 h before next secretion series. A batch of forty-five virginoparae female adults, reproduced by the same virginoparae female adult within 1 h, were introduced onto the tea seedlings (one aphid/seedling) under 13–21°, 85% RH, 3500 lx and 12 L:12 D photoperiod. Temperature showed a significant effect on the amounts of honeydews excreted within the range of 13–21°. Honeydews excreted by the aphids significantly increased the searching and retention time of the tested 9 species of natural enemies in a positive dose-response fashion. The searching times of Aphidius sp. and S. menthastri were the longest and the shortest, respectively, among all the 9 species, while the searching and retention time of L. axyridis (Pallas) var. spectabilis was the longest among the four varieties of L. axyridis. Tea aphid oneydew is considered as an important contact kairomone for the tested natural enemies.     

Red trap colour of the carnivorous plant Drosera rotundifolia does not serve a prey attraction or camouflage function

The traps of many carnivorous plants are red in colour. This has been widely hypothesized to serve a prey attraction function; colour has also been hypothesized to function as camouflage, preventing prey avoidance. We tested these two hypotheses in situ for the carnivorous plant Drosera rotundifolia. We conducted three separate studies: (i) prey attraction to artificial traps to isolate the influence of colour; (ii) prey attraction to artificial traps on artificial backgrounds to control the degree of contrast and (iii) observation of prey capture by D. rotundifolia to determine the effects of colour on prey capture. Prey were not attracted to green traps and were deterred from red traps. There was no evidence that camouflaged traps caught more prey. For D. rotundifolia, there was a relationship between trap colour and prey capture. However, trap colour may be confounded with other leaf traits. Thus, we conclude that for D. rotundifolia, red trap colour does not serve a prey attraction or camouflage function.     

大豆田中大豆蚜天敌昆虫群落结构分析

为了明确大豆田中大豆蚜Aphis glycines Matsumura天敌昆虫的群落结构特点,采用系统调查法对大豆蚜天敌的种类及数量进行调查。结果表明,大豆蚜天敌昆虫主要有5目11科19种,另外还有一些捕食性蜘蛛。主要优势种为小花蝽Orius minutus、龟纹瓢虫Propylaea japonica、中华草蛉Chrysopa sinica以及蚜小蜂Aphelinus sp.和异色瓢虫Leis axyridis,捕食性蜘蛛其中小花蝽的相对丰盛度达到0.26以上。小花蝽和龟纹瓢虫发生较早,是大豆蚜发生初期的主要控制因素。物种多样性、均匀度、丰富度、物种数和个体数整体趋势表现为先增加后降低,在7、8月达最大;而优势集中性和优势度则表现为先降低后增加的趋势,群落较稳定。     

Extra-pair paternity does not result in differential sexual selection in the mutually ornamented black swan (Cygnus atratus)

We studied patterns of parentage in 85 broods (332 cygnets) of black swans during three breeding seasons, using a set of eight polymorphic microsatellite markers. We detected both intraspecific brood parasitism (IBP; < 5% of cygnets per year) and extra-pair paternity (EPP). In these years, 10-17% (mean = 15.1%) of cygnets resulted from EPP, and 27-40% (mean 37.6%) of broods contained at least one extra-pair cygnet. Compared with levels of EPP in closely related species with similar life histories, these values are unexpectedly high. EPP in black swans appears unrelated to ecological factors (breeding density and synchrony) or genetic factors (genetic similarity between pair members or genetic quality of the offspring). We found no evidence that a mutual sexual feather ornament known to play a role in social mate choice in black swans (curled wing feathers) is involved in extra-pair mate choice. EPP does not lead to greater variance in reproductive success in males, relative to females in this species. We therefore suggest that EPP does not result in differential sexual selection on males and females, explaining why they are ornamented to the same degree.     

Heritable variation in the attraction of Drosophila melanogaster to fruit in the field

Drosophila melanogaster were released at a field site and captured with two types of fruit pulp. When flies were rereleased they were more likely to be captured with the same type of fruit on which they were initially captured. Progeny reared in the laboratory tended to be attracted to the same type of fruit as their parents in field tests with two fruit combinations (lemon-orange, lemon-apple) but not a third combination (apple-orange). However, lines selected in the laboratory for increased attraction to apples or oranges over 15 generations differed in their response to these fruit types in the field. This indicates heritable variation for attraction to natural resources under field conditions. Simulations suggest that genotypes differed substantially in their responses to fruit resources.     

茶蚜(Toxoptera aurantii (Boyer))蜜露分泌节律及对多种天敌的引诱效应

为研究茶蚜蜜露的分泌节律及对蚜茧蜂、中华草蛉、大草蛉、门氏食蚜蝇、七星瓢虫、异色瓢虫二斑变型、显现变种、显明变种和十九斑变种的引诱效应,在21℃、85%RH、3500lx光照和12L∶12D光周期下,把1头无翅胎生雌成蚜在1h内产出的45头无翅胎生雌若蚜分别引至小茶苗上,观测发育,收集蜜露。发现若蚜和成蚜历期为32.4d±5.8d,分泌蜜露325.6滴±35.8滴,体积为41.98μl±6.14μl,质量为45.34mg±8.76mg。1～4龄期间的蜜露量与天数呈Logistic关系。其中1～2龄期分泌的蜜露量较小。成蚜历期22.0d±0.0d,分泌蜜露176.31滴±22.38滴,体积30.38μl±5.32μl,连续分泌5～8h之后间歇2～5h。另取一头雌成蚜在1h内产出的、已发育为成蚜的45头雌成蚜,在13～21℃、85%RH、3500lx光照和12L∶12D光周期下观测,发现室温的升降明显影响蜜露分泌量。蜜露显著延长9种(亚种)天敌昆虫搜寻时间;随着蜜露剂量的增大,搜寻时间显著地延长。蚜茧蜂搜寻时间最长,门氏食蚜蝇最短,异色瓢虫4变种(型)中,显明变种搜寻时间最长。茶蚜蜜露是重要的引诱多种天敌的接触性利它素。     

稻田主要天敌对害虫的捕食作用

应用二次正交旋转组合设计方法建立3种天敌、2种害虫共存情况下白背飞虱被捕食数量与5个因素相互关系的数学模型。结果表明,3种天敌对白背飞虱被捕食量影响的大小排序为拟水狼蛛>青翅蚁形隐虫>食虫沟瘤蛛,食虫沟瘤蛛与拟水狼蛛、拟水狼蛛与青翅蚁形隐翅虫有负交互作用。     

Simultaneous feeding by aboveground and belowground herbivores attenuates plant-mediated attraction of their respective natural enemies

Herbivore-damaged plants emit volatile organic compounds that attract natural enemies of the herbivores. This form of indirect plant defence occurs aboveground as well as belowground, but it remains unclear how simultaneous feeding by different herbivores attacking leaves and roots may affect the production of the respective defence signals. We employed a setup that combines trapping of volatile organic signals and simultaneous measurements of the attractiveness of these signals to above and belowground natural enemies. Young maize plants were infested with either the foliar herbivore Spodoptera littoralis , the root herbivore Diabrotica virgifera virgifera , or with both these important pest insects. The parasitic wasp Cotesia marginiventris and the entomopathogenic nematode Heterorhabditis megidis were strongly attracted if their respective host was feeding on a plant, but this attraction was significantly reduced if both herbivores were on a plant. The emission of the principal root attractant was indeed reduced due to double infestation, but this was not evident for the leaf volatiles. The parasitoid showed an ability to learn the differences in odour emissions and increased its response to the odour of a doubly infested plant after experiencing this odour during an encounter with hosts. This first study to measure effects of belowground herbivory on aboveground tritrophic signalling and vice-versa reemphasizes the important role of plants in bridging interactions between spatially distinct components of the ecosystem.     

豇豆田生态系统中主要害虫及天敌的生态位研究

对武汉地区春播和夏播豇豆田生态系统中主要害虫、天敌的种类及空间、时间生态位进行了调查研究。结果表明,春播和夏播豇豆中,瓢虫、蜘蛛和捕食蝽同主要害虫的生态位重叠较大,跟随作用强,具保护利用价值;害虫生态位宽度以蚜虫LipaahiserysimiKaltenbach、蓟马ThripspalmiKarny、豆野螟MarucatestulalisGeyer较宽,天敌生态位春播田中瓢虫较宽,夏播田中蜘蛛较宽。     

中稻田三种飞虱的捕食性天敌优势种及农药对天敌的影响

对两优0923非防治的中稻田白背飞虱、灰飞虱和褐飞虱与其天敌间的关系,采用灰色关联度法、生态位分析方法,对盆拍法调查的3种飞虱与其捕食性天敌在数量、时间和空间三方面关系进行分析,对每一种天敌对应的关联度、生态位重叠指数和相似性比例等参数标准化后的密切指数相加,按照密切指数值之和大小排序,评判3种飞虱捕食性天敌优势种.并用同样方法分析常规防治田农药对飞虱捕食性天敌的影响,以期为合理施药,科学保护和利用天敌优势种提供科学依据,其结果是,非防治田白背飞虱前三位天敌是条纹蝇虎、草间小黑蛛和锥腹肖蛸;灰飞虱的是八斑球腹蛛、茶色新园蛛和锥腹肖蛸;褐飞虱的是纵条蝇狮、四点亮腹蛛和黑肩绿盲蝽.防治田白背飞虱前三位天敌是条纹蝇虎、草间小黑蛛和锥腹肖蛸;灰飞虱的是拟水狼蛛、四点亮腹蛛和草间小黑蛛;褐飞虱的是黑肩绿盲蝽、拟水狼蛛和四点亮腹蛛.盆拍法的防治田和非防治田之间3种飞虱前三位的相同天敌,白背飞虱完全相同,灰飞虱没有相同天敌,褐飞虱的是黑肩绿盲蝽和四点亮腹蛛.飞虱与天敌在时间和数量关系上,扫网法的防治田和非防治田之间3种飞虱前三位的相同天敌,白背飞虱的是锥腹肖蛸和四点亮腹蛛,灰飞虱完全相同,褐飞虱的是纵条蝇狮和条纹影虎.两种稻田的差异主要是农药杀伤了飞虱,使飞虱数量减少,并对天敌有一定杀伤力,进而影响到天敌的发生规律.非防治田的盆拍法和扫网法之间,3种飞虱前三位相同的天敌,白背飞虱的是锥腹肖蛸,灰飞虱的也是锥腹肖蛸,褐飞虱的是纵条蝇狮.防治田两调查方法结果之间,白背飞虱的是锥腹肖蛸,灰飞虱和褐飞虱前3位天敌中没有相同天敌,其差异主要是两法调查稻株的部位不同所致.     

Domestication of tomato has reduced the attraction of herbivore natural enemies to pest‐damaged plants

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