Reactions of adult female parasitoids, particularly Aphidius rhopalosiphi, to volatile chemical cues from the host plants of their aphid prey

Abstract. In Y-tube olfactometer tests, Aphidius ervi Hal., Trioxys sp., Praon sp., Aphelinus flavus (Nees), Lysiphlebus fabarum (Marsh.) and Aphidius rophalosiphi De Stef. responded positively to the odour of the plant on which aphid mummies containing them had been collected. The response to host plant odour was greater than the response to the odour of host aphids, their honeydew or a combination of the two. The strongest response was to a combination of plant and host aphids. A. rhopalosiphi showed a strong positive response to three wheat volatiles (cis-3-hexenyl acetate, cis-3-hexen-1-ol and trans-2-hexenal) as well as to indole-3-acetaldehyde (a breakdown product of tryptophan in aphid honeydew). In both olfactometer tests with odours and choice trials with whole plants, newly emerged A. rhopalosiphi distinguished and preferred the variety of wheat on which their development had occurred to other wheat varieties.     

Aphid parasitoids detect that an alien plant was present nearby during their development

Abstract Glasshouse and laboratory experiments were conducted with Aphidius rhopalosiphi parasitizing Metopolophium dirhodum to test whether the presence in the same environment of another plant species (tomato or marigold) induced aerial allelopathy in wheat plants.
Emerging parasitoids were offered the choice of two odours of wheat plants in an olfactometer – wheat grown alone and wheat grown in the presence of the second plant species.
Female parasitoids responded more strongly than males, but both sexes preferred the odour of wheat grown in the environment in which they had developed (i.e. with or without the second plant species present).
Moving the tomato plants from the cages where they had been placed with wheat to the cages that had previously had wheat alone just before the aphid mummy formed showed that the odour environment the emerging parasitoid responded to was that of the mummy rather than that of the developing parasitoid larva. From this it was concluded that the parasitoids in the olfactometer were responding not only to wheat volatiles, but also to tomato volatiles absorbed on the surface of the mummy and encountered by the emerging adult parasitoid.
By excising the parasitoid pupa from the mummy formed on one wheat cultivar, and offering the emerging parasitoids the surface of aphid mummies formed on a different cultivar, it was confirmed that olfactometer responses represented a learning of the chemistry of the surface of the mummy encountered at or after emergence.     

Falling aphids enhance impact of biological control by parasitoids on partially aphid-resistant plant varieties

Cage experiments, in which the population increase of Metopolophium dirhodum was measured either in the absence or presence of the parasitoid Aphidius rhopalosiphi, showed reductions of peak populations in the presence of the parasitoid of 30% on the susceptible wheat cultivar Armada, but of 57% on the partially resistant Rapier. On both cultivars, aphid population curves in the presence and absence of the parasitoid diverged before the first mummies were formed. The number of aphids leaving plants was measured, and was found to increase in the presence of the parasitoid, particularly with Rapier (almost double). Most aphids (about 75%) which left plants fell, and a smaller proportion regained plants when parasitoids were present than when they were absent. A field experiment with Brevicoryne brassicae and natural predation on Brussels sprouts also showed an increase in aphids leaving a less susceptible compared with a more susceptible cultivar.     

Effects of temperature, feeding position and crop growth stage on the population dynamics of the rose grain aphid, Metopolophium dirhodum (Hemiptera: Aphididae)

The population dynamics of Metopolophium dirhodum were studied on winter wheat seedlings at constant (10°C, 15°C, 20°C, 25°C and 30°C) and fluctuating (12(night)-22(day)°C) temperatures, and during booting to early inflorescence, and anthesis to early milky ripe stage, at 19°C. The pre-reproductive development time was decreased by increasing the temperature from 10°C to 25°C. It was significantly shorter when the aphids were feeding during booting to early inflorescence than during anthesis to early milky ripe stage but was similar when the aphids were feeding on the flag, second or third leaves. The total number of nymphs produced/apterous adult was not significantly affected by temperature from 10°C to 25°C but adult reproductive lifespan was reduced by increasing temperature from 10°C and 12–22°C to 15°C, 20°C and 25°C. The daily intrinsic rate of increase changed from 0.11 to 0.25, and the cohort generation time decreased from 31 to 12 days, with increase of temperature from 10°C to 25°C. Reproductive rate was similar when the aphids were feeding on the flag, second or third leaves during booting to early inflorescence at 19°C. The reproductive rate was higher when the aphids fed from mid-inflorescence to mid-milky ripe stage than from mid-milky to early ripe stage. These results were compared with those from other studies. Predictions from a simulation model using development and reproductive rates from this study and literature were compared and the former rates resulted in a more accurate prediction of field observations in 1979, an outbreak year.     

Plant structure and the foraging success of Aphidius rhopalosiphi (Hymenoptera: Aphidiidae)

Abstract. 1. A study of searching behaviour on wheat and host preference of Aphidius rhopalosiphi (DeStefani-Perez) is described.
2. Parasitoids divided their time equally between the leaves but spent very little time on the ear.
3. After contact with honeydew or an aphid host, parasitoids were arrested in a particular area and increased the time spent searching.
4. Aphids feeding on the ear were parasitized less successfully, since their position between the grains protected them from parasitoid attacks.
5. A.rhopalosiphi exhibited no preference for Metopolophium dirhodum (Walker) or Sitobion avenae (Fabricius), although the handling time for the latter was significantly longer; this resulted in fewer S.avenae being parasitized when it was abundant. Parasitoids did not switch between hosts in these experiments.
6. As a result of its searching behaviour, A.rhopalosiphi will encounter and parasitize M.dirhodum feeding on the leaves more frequently than S.avenae, which feeds on the ear. This will limit the parasitoid's ability to regulate populations of the cereal aphid S. avenae.     

A maternal influence on the conditioning to plant cues of Aphidius colemani Viereck, parasitizing the aphid Myzus persicae Sulzer

Abstract. The offspring of parasitoids, Aphidius colemani Viereck, reared on Brussels sprouts and emerging from Myzus persicae Sulzer on a fully defined artificial diet, show no preferences in a four-way olfactometer, either for the odour of the diet, the odour of Brussels sprouts, or the odour of two other crucifers (cabbage and Chinese cabbage). A similar lack of odour preferences is shown when the host aphids are exposed for parasitization (for 48 h) on cabbage, Chinese cabbage or wheat. However, if parasitization occurs on Brussels sprouts, a weak but statistically highly significant response to Brussels sprout odour is observed. Although as many as 30–35% of the parasitoids show no response to any odour, another 35% respond positively to the odour of Brussels sprout compared with responses to the odours of cabbage, Chinese cabbage or wheat of only approximately 10%. An analagous result is obtained when the parent parasitoids are reared on cabbage. In this case, significant positive responses of their offspring to cabbage odour occur only if the 48-h parasitization has occurred also on cabbage. However, with parasitoids from Brussels sprouts parasitizing the aphids for 48 h also on Brussels sprouts, the offspring subsequently emerging from pupae excised from the mummies show no preference for Brussels sprout odour. Thus, although the Brussels sprout cue had been experienced early in the development of the parasitoids, they only become conditioned to it when emerging from the mummy. Both male and female parasitoids respond very similarly in all experiments. It is proposed that the chemical cue (probably glucosinolates in these experiments) is most likely in the silk surrounding the parasitoid pupa, and that the mother may leave the chemical in or around the egg at oviposition, inducing chemical defences in her offspring to the secondary plant compounds that the offspring are likely to encounter.     

The influence of the host plant of diamond-back moth (Plutella xylostella) on the plant preferences of its parasitoid Cotesia plufellae in Sri Lanka

Abstract. In Y-tube olfactometer tests, female Cotesia plutellae Kurdjmov (Hymenoptera: Braconidae), a major parasitoid of Plutella xylostella (L.) in southeast Asia, preferred the odour of the brassica type (Blue Lake cabbage) on which they had developed.Such plants were even preferred to the odour of caterpillar-infested and damaged leaves of another brassica (Early Phenomenal cauliflower).However, within one brassica type, the odour of caterpillar-infested leaves was preferred to that of uninfested and undamaged leaves.Odours of both Blue Lake and Early Phenomenal were preferred to the control odour (water).However, following the transfer of the parasitoid from Blue Lake to another cabbage (Brunswick) for one generation, females no longer distinguished between Blue Lake and Early Phenomenal.     

Abundance of the rose-grain aphid,Metopolophium dirhodum,on barley in Canterbury,New Zealand, 1984–87

Abstract

Spring populations of Metopolophium dirhodum (Walker) (Hemiptera: Aphididae) on barley, at Lincoln and elsewhere in Canterbury, New Zealand, declined markedly during 1984–87. Spring suction-trap catches of M. dirhodum also declined at the same time. Numbers of immigrant M. dirhodum on young barley, and maximum aphid population density on the crop, were positively correlated with suction-trap catches of M. dirhodum during the preceding week or month, respectively. Egg numbers of the most abundant predator, Micromus tasmaniae Walker (Neuroptera: Hemerobiidae), and mummy numbers of the parasitoid, Aphidius rhopalosiphi De Stefani-Perez (Hymenoptera: Aphidiidae), increased relative to aphid numbers during the study. Entomophagous fungus-disease incidence could not be related to monthly rainfall, and the decline in aphid numbers during the study period was apparently not related to disease incidence. It is suggested that aphid population regulation by predators and parasitoids in the wider habitat of M. dirhodum could reduce the rate of immigration, and hence aphid population growth, in a given crop.     

Factors determining the pest status of the rose-grain aphid, Metopolophium dirhodum (Walker), on winter barley in the United Kingdom

To test the hypothesis that early maturation of barley affords it some resistance to cereal aphids, the colonisation, survival, growth, reproduction and emigration of M. dirhodum on barley was measured and compared with previously published results for this aphid on wheat.
Barley plants were colonised as readily as wheat plants. Survival to maturity was similar on both hosts. Relative growth rate was higher on wheat than on barley for most of the season while reproduction was similar on both. Emigration, measured alate production, was significantly higher on mature wheat than on mature barley.
It is proposed that the increase of M. dirhodum populations on barley is normally terminated early in the year directly plant maturation and consequent drying out of the leaves, resulting in the death of aphids. This is different to the situation on wheat, which matures later, and where emigration following plant induced alate induction has been shown to be the major factor affecting the decline in population growth.
These results support the hypothesis that early maturation of barley provides some resistance to M. dirhodum.     

The relationship between host selection behaviour and offspring fitness in a koinobiont parasitoid

1. When host quality varies, optimal foraging theory assumes that parasitic wasps select hosts in a manner that increases their individual fitness. In koinobiont parasitoids, where the hosts continue developing for a certain period of time after parasitisation, host selection may not reflect current host quality but may be based on an assessment of future growth rates and resources available for the developing larvae. 2. When presented with hosts of uniform quality, the koinobiont parasitoid Leptomastix dactylopii exhibits a characteristic host‐selection behaviour: some hosts are accepted for oviposition on first encounter, while others are rejected several times before an egg is laid in them, a behaviour that is commonly associated with a changing host acceptance threshold during the course of a foraging bout. 3. The fitness of the offspring that emerged from hosts accepted immediately upon encounter was compared with the fitness of offspring emerged from hosts rejected several times before being accepted for oviposition. 4. The pattern of host acceptance and rejection was not related to any of the measured fitness parameters of the offspring emerging from these hosts (development time, size at emergence, sex ratio at emergence, and female offspring egg load). 5. While complex post facto adaptive explanations can be devised to explain the nature of such a time and energy consuming host selection process, it is suggested that physiological constraints on egg production or oviposition may provide an alternative, purely mechanistic, explanation for the results obtained.     

寄主挥发物、叶色和表皮毛在美洲斑潜蝇寄主选择中的作用

在室内条件下 ,初步研究了寄主挥发物、叶色和表皮毛在美洲斑潜蝇寄主选择中的作用。在嗅觉仪试验中 ,寄主叶片挥发物对美洲斑潜蝇雌成虫没有明显的引诱作用 ;在叶色反应试验中 ,美洲斑潜蝇雌成虫在叶子圆片上停留的时间明显大于在滤纸圆片上停留的时间 (p<0 .0 1) ,其在有叶片区域分布的数量明显多于空白对照 (p <0 .0 1) ;在表皮毛试验中 ,美洲斑潜蝇在无毛叶片上的产卵量明显大于在有毛叶片上的产卵量 (p <0 .0 1)。上述结果表明 ,在对寄主的定向和定位过程中 ,美洲斑潜蝇的视觉起着重要的作用 ,而嗅觉不起作用 ;叶片表皮毛有抑制产卵的作用     

Role of olfaction in host plant selection and local adaptation of a polyphagous herbivore,Eucolaspis Sharp

Host plant cues are known to shape insect–host plant association in many insect groups. More pronounced associations are generally manifested in specialist herbivores, but little is known in generalist herbivores. We used a polyphagous native beetle from New Zealand, bronze beetle, Eucolaspis sp. ‘Hawkes Bay’ (Chrysomelidae: Eumolpinae) to explore the role of olfaction in locating host plants and local adaptation. We also tested the role of other cues in the degree of acceptance or rejection of hosts. Adult Eucolaspis beetles were attracted to fresh leaf volatiles from apple and blackberry (Rosaceae). Male and female beetles responded similarly to olfactory cues of host plants. An indication of evolutionary affiliation was observed in olfactory preferences of geographically isolated conspecific populations. We found that geographically isolated populations of the beetles differ in their olfactory responses and exhibit some degree of local adaptation. However, irrespective of geographical and ecological associations, blackberry was preferred over apple as a feeding plant, and another novel plant, bush lawyer (Rubus australis), was readily accepted by 53.25% of the tested beetles. We show that plant volatiles play an important role in host location by Eucolaspis, but the acceptance or rejection of a particular host could also involve visual and contact cues.     

Leaf buds, a factor in host selection by Battus philenor butterflies

ABSTRACT.

• 1 Field and laboratory experiments identified a character intrinsic to Aristolochia reticulata Nutt. host plants, the terminal leaf bud, that is involved in host-selection behaviour by female pipevine swallowtail butterflies (Battus philenor L.) searching for oviposition sites.
• 2 In the field, the frequency with which females landed on non-host buds declined seasonally as the proportion of host foliage that consisted of buds decreased. Female butterflies did not land on non-host species in proportion to their abundance; rather, females landed on those non-host species whose buds resembled those of A.reticulata.
• 3 A.reticulata plants whose terminal leaf bud was concealed by plastic tape were less susceptible to oviposition in the field than were control plants.
• 4 Female butterflies released in a large, outdoor enclosure were conditioned to search for leaf buds only when exposed to a host species bearing a prominent terminal leaf bud.
• 5 The significance of conditioning of leaf-bud searching behaviour is discussed with respect to discrimination between hosts and non-hosts, between host species, and among plants within a host species.

     

Host plant selection behaviour of Chilo partellus and its implication for effectiveness of a trap crop

Female lepidopterans can display a hierarchy of preference among potential host species, a trait thought to arise from the balance between attractants and deterrents to which the insects respond. Host plant ranking by moths and larvae of Chilo partellus Swinhoe (Lepidoptera: Crambidae), an important pest of cereals in Africa, was investigated, and whether eggs deposited on specific host plants yield larvae of particular host preferences. Trap plants are used in management of this pest. However, any ‘disagreement’ in host ranking between moths and larvae could potentially reduce effectiveness of trap crops as larvae emigrate to the main crop from the parent’s preferred trap plant. We also investigated whether host plant preference is influenced by the diet upon which larvae fed as part of an integrated assessment of the relationship between host plant selection and learning in C. partellus. Five host plants (all Poaceae) were used: maize (Zea mays L.), sorghum (Sorghum bicolor Moench), Napier grass (Pennisetum purpureum Schumach), and two varieties of signal grass [Brachiaria brizantha (A. Rich.) Stapf], viz., local (henceforth signal grass) and improved (‘Mulato’). In multiple choice tests, C. partellus female moths preferentially oviposited on Napier grass, followed by sorghum, maize, and signal grass, and least preferred ‘Mulato’. Larvae however equally orientated and settled on leaf cuts of maize, sorghum, signal grass, and Napier grass, but least preferred ‘Mulato’. Moreover, eggs from specific host plants did not yield larvae of particular host preferences. Furthermore, oviposition preference was not altered by the larval food. These results imply only a slight ‘disagreement’ in host ranking behaviour between moths and larvae, which is beneficial for trap cropping as larvae would not ‘reject’ the trap plant and appreciably disperse to the neighboring plants. Moreover, absence of larval learning behaviour indicates that regardless of the larval food C. partellus moths would still be attracted to the selected trap plant.     

Relationship between volatile compounds of Picea likiangensis var. linzhiensis cone and host selection of Dioryctria abietella

This study aimed to determine the relationship between volatile compounds of Picea likiangensis var. linzhiensis cone and host selection of Dioryctria abietella. During the infestation of P. likiangensis var. linzhiensis by D. abietella, their cones and branches emitted volatile compounds, which were extracted using CH₂Cl₂ extraction and XAD₂ adsorption methods, and were analyzed using gas chromatography–mass spectrometry. Before and after overwintering, D. abietella larva preferred annually infested cones and their extracts, and adult D. abietella preferred to lay eggs on annually infested cones and healthy cones of the year, and the oviposition rate of adult D. abietella was 72% on branches with healthy cones of the year, and no egg was laid on branches with annually healthy cones or branches without cones. The volatile compounds after infestation, α- and β-pinene, were significantly higher in cones than those in other tissues; however, myrcene in cones was significantly lower than those in other tissues. The annually infested cones produced β-caryophyllene and (1S)-(-)-β-pinene, while the annually healthy cones and branches produced myrcene and 3-carene. The annually infested cones and their extracts attracted D. abietella larvae, while that of healthy cones and annually infested cones attracted the adults, indicating that the terpene compounds: α-pinene, β-pinene, (1S)-(-)-β-pinene, limonene, and β-caryophyllene are attractive to D. abietella, and the terpene compounds—myrcene and 3-carene—from the branch tissues may be repulsive to D. abietella.     

Temperature influences host instar selection in an aphid parasitoid: support for the relative fitness rule

The relative fitness rule states that parasitoid females should adopt risk‐prone reproductive behaviours when expecting low reproductive success. Temperature influences the reproductive success of insects by affecting their basal metabolic rate during development, their egg load at emergence, and their life expectancy as adults. Using an aphid–parasitoid model system, we investigated the influence of developmental and adult temperature on the risk‐sensitive decision‐making of females. We considered the use of a low‐quality host nymphal instar by the aphid parasitoid Aphidius ervi to be a risk‐prone behaviour. Immature females were reared at 12, 20 or 28 °C and had access to the four nymphal instars of the potato aphid Macrosiphum euphorbiae for oviposition at one of these temperatures. Host selection behaviour was continuously recorded during exploitation of an aphid patch. We observed that warm‐developed females and parasitoids foraging at high temperature attacked low‐quality hosts more frequently than females from other treatments. These results support the hypothesis that a decrease in expected parasitoid reproductive success resulted in risk‐prone behaviours. To our knowledge, this is the first experimental evidence suggesting that temperature influences host stage selection and risk‐sensitive making decision in parasitoids, and the present study is the first to support the relative fitness rule. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●●, ●●–●●.     

植食性昆虫的嗅觉选食过程及其机制研究进展

植食性昆虫的嗅觉在其选择食物的过程中发挥了重要的作用,它能通过对植物挥发物的感受来定向和定位食物源并产生趋近行为,进而根据特殊的化合物或者多种化合物的特异浓度组合来区分寄主和非寄主植物.在这个过程中,昆虫嗅觉器官上相关的嗅觉感受蛋白被植物挥发物激活,形成特异的嗅觉感受通路,在行为上调控昆虫嗅觉选食的能力.本文主要从植食...     

寄主植物-蚜虫-天敌三重营养关系的化学生态学研究进展

综述了寄主植物－蚜虫－天敌三重营养关系的化学生态学研究,重点阐述了3个研究热点：①植物挥发性物质在蚜虫及其天敌选择寄主行为过程中的作用;②蚜虫信息素和蜜露对蚜虫天敌寄主选择行为的影响;③植物挥发性物质对蚜虫信息系作用的影响。对寄主植物－蚜虫－天敌三重营养关系的全面了解,将为蚜虫的综合治疗提供新思维。     

Response of the pine engraver, Ips pini (Say) (Coleoptera: Scolytidae), to conophthorin and other angiosperm bark volatiles in the avoidance of non-hosts

1 Seventeen non‐host angiosperm bark volatiles, seven of which are antennally active to Ips pini (Say), the pine engraver (PE), were tested for their ability to disrupt the response of the PE to pheromone‐baited traps. 2 Four green leaf volatiles (GLVs) were tested [1‐hexanol (Z)‐3‐hexen‐1‐ol, hexanal, and (E)‐2‐hexenal]. None had any disruptive effect singly, as a group or in all possible blends based on functional groups, despite the fact that the two aldehydes were antennally active. These compounds may have, in some instances, actually masked the disruptive effect of other compounds. The PE thus differs in its response from other Scolytidae, including other Ips spp. 3 Eight non‐host volatiles that were antennally active to other bark beetles, but not to PEs, had no disruptive effect, validating the use of coupled gas chromatographic‐electroantennographic detection analyses to detect compounds with potential behavioural activity. 4 The bicyclic spiroacetal conophthorin, (E)‐7‐methyl‐1,6‐dioxaspiro[4.5]decane, was disruptive when tested alone. When blends of two aldehydes [salicylaldehyde and nonanal] plus an alcohol and a phenol [benzyl alcohol and guaiacol] were combined with conophthorin, an enhanced disruptive effect was revealed. No single compound, other than conophthorin, disrupted the pheromone‐positive response and no blend that did not contain conophthorin was consistently disruptive to both sexes. Conophthorin seems to be a critical component in the non‐host angiosperm message for I. pini during its host selection phase. 5 Combination of the repellent synomones, verbenone and ipsenol, with the five disruptive non‐host volatiles may provide a potent treatment to protect trees, logs or stands from attack by the PE.     

Oviposition site selection by herbivorous beetles: a comparison of two thistle feeders, Cassida rubiginosa and Henosepilachna niponica

In insects that feed on plants in both adult and larval stages, it is often difficult to distinguish oviposition preference from adult feeding preference, because oviposition can occur at or in proximity to feeding sites. In the present study, characteristics of oviposition site selection of two beetle species, Cassida rubiginosa Müller (Coleoptera: Chrysomelidae) and Henosepilachna niponica (Lewis) (Coleoptera: Coccinellidae), were investigated in the field and laboratory, with particular attention to relationships with adult feeding sites. In the field, distances between adult feeding scars and egg masses differed for C . rubiginosa and H . niponica , with the former being very small and the latter averaging 24.6 cm. The same tendencies for the distances between adult feeding scars and egg masses of the two beetle species were confirmed in cages in which only female beetles were released. Cassida rubiginosa restricted egg laying to host plants in the field and to leaves in laboratory assays. On the other hand, H . niponica placed 8% of egg masses on plants adjacent to host plants in the field and often placed eggs on artificial substrates rather than leaf discs in laboratory assays. These results suggest that oviposition and female feeding sites are virtually inseparable in the case of C . rubiginosa , while H . niponica females do not necessarily keep to host plant leaves as oviposition substrates and they tend to oviposit at some distance from their feeding sites. Results are discussed in relation to proximate and ultimate causes of host selection behavior.