Olfactory response of predatory mites to vegetative and reproductive parts of coconut palm infested by <Emphasis Type="Italic">Aceria guerreronis</Emphasis>

The phytophagous mite Aceria guerreronis Keifer is an important pest of coconut worldwide. A promising method of control for this pest is the use of predatory mites. Neoseiulus baraki (Athias-Henriot) and Proctolaelaps bickleyi Bram are predatory mites found in association with A. guerreronis in the field. To understand how these predators respond to olfactory cues from A. guerreronis and its host plant, the foraging behavior of the predatory mites was investigated in a Y-tube olfactometer and on T-shaped arenas. The predators were subjected to choose in an olfactometer: (1) isolated parts (leaflet, spikelet or fruit) of infested coconut plant or clean air stream; (2) isolated parts of non-infested or infested coconut plant; and (3) two different plant parts previously shown to be attractive. Using T-shaped arenas the predators were offered all possible binary combinations of discs of coconut fruit epidermis infested with A. guerreronis, non-infested discs or coconut pollen. The results showed that both predators were preferred (the volatile cues from) the infested plant parts over clean air. When subjected to odours from different infested or non-infested plant parts, predators preferred the infested parts. Among the infested plant parts, the spikelets induced the greatest attraction to predators. On the arenas, both predators preferred discs of coconut fruits infested with A. guerreronis over every other alternative. The results show that both predators are able to locate A. guerreronis by olfactory stimuli. Foraging strategies and implications for biological control are discussed.     

Volatile spider-mite pheromone and host-plant kairomone, involved in spaced-out gregariousness in the spider mite Tetranychus urticae

ABSTRACT. From the host plant-spider mite complex Phaseolus lunatus—Tetranychus urticae Koch a volatile chemical is emitted that acts as a kairomone for the predatory mite Phytoseiulus persimilis Athias-Henriot (Sabelis et al. , 1984a). This kairomone is apparently a byproduct of a vital physiological process and/or it has a function in the biology of the spider mite as well.
The spider mite—host plant complex also emits a volatile spider-mite dispersing pheromone. This is shown in the present study where spider mites were introduced into an odour patch on a horizontal screen in a vertical airflow olfactometer. When spider-mite infested leaves of Lima bean are offered, the spider mites walk mainly straight and soon reach the edge of the screen. On the other hand, when clean Lima bean leaves are offered, the mites walk tortuously most of the time and reach the edge of the screen much later. Artificially damaged plants elicit the same response as undamaged plants. Differences in spider-mite behaviour are observed in the vertical airflow olfactometer when odour of either clean or spider-mite infested leaves is offered. A comparison of the behaviour in these two situations with that when no odour was offered suggests that Lima bean leaves emit a volatile kairomone that activates T. urticae and makes them return after losing the stimulus. A Y-tube olfactometer experiment confirms the existence of this kairomone.
At a low ratio of dispersing pheromone to plant kairomone, the spider mites behave as if only kairomone is present, walking mainly tortuously. At a high ratio they disperse. No aggregation-pheromonal effect is observed.
The possibility that the spider-mite dispersing pheromone acts as a kairomone for P. persimilis is discussed.     

Host race formation inTetranychus urticae: genetic differentiation, host plant preference, and mate choice in a tomato and a cucumber strain

The two-spotted spider mite,Tetranychus urticae Koch, occurs in two colour forms in greenhouses in the Netherlands: a red form on tomato and a green form on cucumber. The evolutionary status of these strains was analysed by studying genetic differentiation, host plant preference, and mate choice. Males of the tomato strain preferred the female (30 h-old) teleiochrysales from the same strain to those of the cucumber strain, independent of the host plants (tomato, cucumber, bean) on which the teleiochrysales were placed. In contrast, males of the cucumber strain were not selective. In a Y-tube olfactometer, females of the cucumber strain were not responsive to host plant volatiles alone. However, in two-choice disc experiments, where females were exposed to both volatile and contact cues, they settled on cucumber leaves in preference to tomato leaves. Females of the tomato strain preferred the odour of tomato leaves and settled on tomato leaves in preference to cucumber leaves. These experimental results provide the first evidence for (1) host-plant independent mate selection in male spider mites and (2) olfactory discrimination between host plants in female spider mites. Electrophoretic analysis showed much genetic differentiation at the phosphoglucose isomerase locus. The cucumber strain showed large variation with 5 alleles, whereas the tomato strain was fixed for the most common allele of the cucumber strain. The results suggest that the two strains represent host races. We hypothesize that the tomato strain has originated from the cucumber strain because (1) tomato represents a more hostile host plant to spider mites (due to toxic compounds and glandular hairs) and (2) the tomato strain is genetically impoverished suggesting that it passed through one or more bottlenecks.     

Volatile and Contact Chemicals Released by Nezara viridula (Heteroptera:Pentatomidae) Have a Kairomonal Effect on the Egg Parasitoid Trissolcus basalis (Hymenoptera: Scelionidae)

The responses of females of the egg parasitoid Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae) to volatile and contact chemicals from its host Nezara viridula (L.) (Heteroptera: Pentatomidae) were investigated in a Y-tube olfactometer and under open arena conditions. In the Y-tube tests, volatiles from virgin males and from females in a preovipositional state attracted T. basalis females, while volatiles from host virgin females did not. In an open arena, traces left by N. viridula adults in different physiological conditions function as contact cues inducing the wasps to remain longer in the arena and to change the pattern of their walking behavior. However, only contact kairomones from N. viridula mated females in a preovipositional condition induced an arrestment response characterized by an increase in patch searching time and turning rates and a reduction in linear speed. The chemical ecological implications of these results on this host–parasitoid association are discussed.     

Response of predatory mites with different rearing histories to volatiles of uninfested plants

The behavioural response of the predatory mite Phytoseiulus persimilis to volatiles from several host plants of its prey, spider mites in the genus Tetranychus, was investigated in a Y-tube olfactometer. A positive response to volatiles from tomato leaves and Lima bean leaves was recorded, whereas no response was observed to volatiles from cucumber leaves, or leaves of Solanum luteum and Solanum dulcamara.Different results were obtained for predators that differed in rearing history. Predators that were reared on spider mites (Tetranychus urticae) on Lima bean leaves did respond to volatiles from Lima bean leaves, while predators that had been reared on the same spider mite species but with cucumber as host plant did not respond to Lima bean leaf volatiles. This effect is compared with the effect of rearing history on the response of P. persimilis to volatile allelochemicals of prey-infested plant leaves.     

The role of plant chemical cues in determining host preference in the generalist aphid parasitoid Aphidius colemani

The host preference behaviour of the generalist aphid parasitoid Aphidius colemaniwas investigated using a Y-tube olfactometer. Female A. colemanishowed a preference for the host-plant complex on which they had been reared, even though the same aphid host was involved, demonstrating a host plant preference. This preference was not evident when the parasitoids were dissected from their mummies prior to adult emergence. Host plant preference exhibited during host selection appeared to be induced by chemical cues encountered on the mummy case at the time of emergence, but preferences could be changed by subsequent foraging experiences. It is concluded that plant chemical cues play a major role in determining initial preferences through a process of emergence conditioning but that learning processes, involving cues encountered during oviposition in or contact with the host, can modify these initial preferences.     

The role of two plant-derived volatiles in the foraging movement of 1st instar Helicoverpa armigera (Hübner): time to stop and smell the flowers

Positive phototaxis and negative geotaxis are behaviours that 1st instar Helicoverpa armigera use to direct their foraging movement upward towards nutritious new plant growth and reproductive structures. Odours emitted by fruits or seeds can attract larvae directly via chemotaxis. In this study we clarify the effect of leaf and flower odours on foraging 1st instar H. armigera. Using a Y-tube olfactometer we tested for chemotaxis towards two plant volatiles and found larvae were not attracted. Bioassays for phototaxis towards UV, blue, green and white light showed that a green leaf volatile ((Z)-3-hexenyl acetate) and a flower volatile (phenylacetaldehyde) reduced larval phototaxis towards blue light. Feeding on a host plant reduced phototaxis towards blue and green light. We concluded that the upward movement of 1st instars on plants is largely due to phototaxis towards the blue wavelengths of skylight. Plant attributes such as volatile chemicals affect the expression of phototaxis and therefore, indirectly influence larval movement to locate food resources. Handling Editor: Anna-Karin Borg-Karlson     

Response of a phytoseiid predator to herbivore-Induced plant volatiles: Selection on attraction and effect on prey exploitation

Bean plants infested with herbivorous spider mites emit volatile chemicals that are attractive toP. persimilis, a predator of spider mites. In Y-tube olfactometer tests we evaluated involvement of a genetic component in predator response to herbivore-induced plant volatiles. Replicated bidirectional selection resulted in a significant increase in attraction after one generation of selection, but no decrease even after three generations of selection, indicating significant, but unbalanced, additive genetic variation in predator perception of, or response to, herbivore-induced plant volatiles. Selected lines responded differently than an unselected population to food deprivation, pointing to an interaction between their internal state and response to plant volatiles. Selected lines also differed from unselected ones in behaviors associated with local prey exploitation, such as residence time, prey consumption, and reproduction. At lower prey densities,P. persimilis from both “+” lines left spider mite-infested leaves more rapidly and consumed fewer prey eggs than an unselected population. Defining olfactory components of predator search behavior is one step in understanding the effect of plant volatiles on predator foraging efficiency. By selecting lines differing in their attraction to herbivore-induced plant volatiles we may experimentally investigate the link between this behavior, predator foraging efficiency, and local and regional predator-prey population dynamics. The impact of significant additive genetic variation in predator response to plant volatiles on evolution in a tritrophic context also remains to be uncovered.     

Limits to ambulatory displacement of coconut mites in absence and presence of food-related cues

Ambulatory movement of plant-feeding mites sets limits to the distances they can cover to reach a new food source. In absence of food-related cues these limits are determined by survival, walking activity, walking path tortuosity and walking speed, whereas in presence of food the limits are also determined by the ability to orient and direct the path towards the food source location. For eriophyoid mites such limits are even more severe because they are among the smallest mites on earth, because they have only two pairs of legs and because they are very sensitive to desiccation. In this article we test how coconut mites (Aceria guerreronis Keifer) are constrained in their effective displacement by their ability to survive in absence of food (meristematic tissue under the coconut perianth) and by their ability to walk and orient in absence or presence of food-related cues. We found that the mean survival time decreased with increasing temperature and decreasing humidity. Under climatic conditions representative for the Tropics (27 °C and 75 % relative humidity) coconut mites survived on average for 11 h and covered 0.4 m, representing the effective linear displacement away from the origin. Within a period of 5 h, coconut mites collected from old fruits outside the perianth moved further away from the origin than mites collected under the perianth of young fruits. However, in the presence of food-related cues coconut mites traveled over 30 % larger distances than in absence of these cues. These results show that ambulatory movement of eriophyoid mites may well bring them to other coconuts within the same bunch and perhaps also to other bunches on the same coconut palm, but it is unlikely to help them move from palm to palm, given that palms usually do not touch each other.     

The influence of conditioning on olfactory microhabitat and host location in Asobara tabida (Nees) and A. rufescens (Foerster) (Braconidae: Alysiinae) larval parasitoids of Drosophilidae

Olfactory responses of parasitoids can be variable. This was shown by olfactometer experiments with females of two sibling Asobara species, larval endoparasitoids of Drosophilidae. Oviposition experiences of adult female parasitoids significantly altered their behavioural responses in microhabitat and host location. Females needed prior exposure to a host before they were capable of using volatile compounds related to the presence of their host. Asobara tabida (Nees) attacks Drosophila in fermenting fruits and A. rufescens (Foerster) attacks Drosophilidae in decaying plant materials. Naïve females show a strong preference for the odour of their own microhabitat. After experience with their own host and microhabitat, females were repelled by odours of the other microhabitat. Enforced experience with this repellent microhabitat in the laboratory modified the olfactory response from repellency to attraction. It was shown that even the microhabitat odour preference pattern could be changed through experience. This kind of behavioural flexibility may be the rule rather than the exception in many other Hymenopterous parasitoids.     

捕食螨化学生态研究进展

捕食螨是重要的生物防治因子。早在20世纪70年代就发现了捕食螨的性信息素,许多研究证明植物挥发物在捕食螨向猎物定位过程中发挥着至关重要的作用,影响捕食螨寻找猎物的植物挥发物来源于未受害植物、机械损伤植物、猎物危害植物、非猎物危害植物。人工合成的植物挥发物组分对捕食螨具有引诱作用,但引诱活性低于虫害诱导植物释放的挥发性混合物。捕食螨的饲养条件、饥饿程度、学习与经验行为等会影响捕食螨对植物挥发物的反应。介绍了信息素与植物挥发物对捕食螨的作用,并讨论了目前存在的问题和研究前景。     

Below-ground plant parts emit herbivore-induced volatiles: olfactory responses of a predatory mite to tulip bulbs infested by rust mites

Although odour-mediated interactions among plants, spider mites and predatory mites have been extensively studied above-ground, belowground studies are in their infancy. In this paper, we investigate whether feeding by rust mites (Aceria tulipae) cause tulip bulbs to produce odours that attract predatory mites (Neoseiulus cucumeris). Since our aim was to demonstrate such odours and not their relevance under soil conditions, the experiments were carried out using a classic Y-tube olfactometer in which the predators moved on a Y-shaped wire in open air. We found that food-deprived female predators can discriminate between odours from infested bulbs and odours from uninfested bulbs or artificially wounded bulbs. No significant difference in attractiveness to predators was found between clean bulbs and bulbs either wounded 30 min or 3 h before the experiment. These results indicate that it may not be simply the wounding of the bulbs, but rather the feeding by rust mites, which causes the bulb to release odours that attract N. cucumeris. Since bulbs are belowground plant structures, the olfactometer results demonstrate the potential for odour-mediated interactions in the soil. However, their importance in the actual soil medium remains to be demonstrated.     

Spider mites adaptively learn recognizing mycorrhiza-induced changes in host plant volatiles

Symbiotic root micro-organisms such as arbuscular mycorrhizal fungi commonly change morphological, physiological and biochemical traits of their host plants and may thus influence the interaction of aboveground plant parts with herbivores and their natural enemies. While quite a few studies tested the effects of mycorrhiza on life history traits, such as growth, development and reproduction, of aboveground herbivores, information on possible effects of mycorrhiza on host plant choice of herbivores via constitutive and/or induced plant volatiles is lacking. Here we assessed whether symbiosis of the mycorrhizal fungus Glomus mosseae with common bean plants Phaseolus vulgaris influences the response of the two-spotted spider mite Tetranychus urticae to volatiles of plants that were clean or infested with spider mites. Mycorrhiza-naïve and -experienced spider mites, reared on mycorrhizal or non-mycorrhizal bean plants for several days before the experiments, were subjected to Y-tube olfactometer choice tests. Experienced but not naïve spider mites distinguished between constitutive volatiles of clean non-mycorrhizal and mycorrhizal plants, preferring the latter. Neither naïve nor experienced spider mites distinguished between spider mite-induced volatiles of mycorrhizal and non-mycorrhizal plants. Learning the odor of clean mycorrhizal plants, resulting in a subsequent preference for these odors, is adaptive because mycorrhizal plants are more favorable host plants for fitness of the spider mites than are non-mycorrhizal plants.     

Lotus japonicus Infested with Herbivorous Mites Emits Volatile Compounds That Attract Predatory Mites

Lotus Japonicus has an indirect defense mechanism against spider mites, Tetranychus urticae, we investigated the responses of predatory mites, Phytoseiulus persimilis, to volatile compounds released from T. urticae-infested L. japonicus in a Y-tube olfactometer. Plants infested with spider mites attracted more P. persimilis than did clean air. Uninfested plants and artificially damaged plants did not attract P. persimilis. When infested by spider mites, L. japonicus plants started emitting (Z)-3-hexenyl acetate, (E)-4,8-dimethyl-1,3,7-nonatriene, germacrene d, 1-octen-3-ol and methyl salicylate (MeSA). These compounds were considered to be T. urticae-induced plant volatile compounds. When three L. japonicus mutants deficient in nodule organogenesis were infested by the spider mites, they all attracted P. persimilis. However, two of the infested mutants emitted blends of induced volatile compounds that were qualitatively different from those emitted from infested wild type L. japonicus. Received 8 August 2000/ Accepted in revised form 12 October 2000     

OLFACTORY RESPONSES OF LYSIPHLEBIA JAPONICA TO VOLATILE CHEMICALS AND FRESH LEAVES OF THE HOST PLANTS OF COTTON APHIDS IN OLFACTOMETER

Abstract The behavioral responses of the female adult parasitoids Lysiphlebia japonica to plant volatile chemicals, host plants and cotton aphids were tested in a four arm air flow olfactometer. The results indicated that the female parasitoids were strongly attracted by the odors from cotton aphids and their host plants. Damaged cotton leaves were more attractive than intact ones. Among the four olfactory stimuli from the plant volatile chemicals dissolved in liquid paraffin oil at the concentration of 0.01 (volume fraction), the female parasitoids only positively responded to benzaldehyde, but did not show any preference to β-caryophyllene and β-pinene. Although heptanol elicited the highest EAC response of the parasitoids, the behavioral test showed that it repelled the parasitoids, and β-pinene shared the same feature with heptanol at the concentration of 1 (volume fraction). Our results also showed that the female parasitoids only preferred the cotton leaf from which the mummified aphids were collected, and did not express any interest to the three plants of melon family in the olfactometer, The possible mechanisms are discussed.     

Can chemical signals,responsible for mutualistic partner encounter,promote the specific exploitation of nursery pollination mutualisms? – The case of figs and fig wasps

In nursery pollination mutualisms, where pollinators reproduce within the inflorescence they pollinate, floral scents often play a major role in advertizing host location and rewards for the pollinator. However, chemical messages emitted by the plant that are responsible for the encounter of mutualist partners can also be used by parasites of these mutualisms to locate their host. Each species of Ficus (Moraceae) is involved in an obligatory nursery pollination mutualism with usually one pollinating fig wasp (Hymenoptera: Chalcidoidea: Agaonidae). In this interaction, volatile compounds emitted by receptive figs are responsible for the attraction of their specific pollinator. However, a large and diverse community of non-pollinating chalcidoid wasps can also parasitize this mutualism. We investigated whether the chemical message emitted by figs to attract their pollinator can promote the host specificity of non-pollinating fig wasps. We analysed the volatile compounds emitted by receptive figs of three sympatric Ficus species, namely, Ficus hispida L., Ficus racemosa L., and Ficus tinctoria G. Forster, and tested the attraction of the pollinator of F. hispida ( Ceratosolen solmsi marchali Mayr), and of one species of non-pollinating fig wasp [ Philotrypesis pilosa Mayr (Hymenoptera: Chalcidoidea: Pteromalidae)] to scents emitted by receptive figs of these three Ficus species. Analysis of the volatile compounds emitted by receptive figs revealed that the three Ficus species could be clearly distinguished by their chemical composition. Behavioural bioassays performed in a Y-tube olfactometer showed that both pollinator and parasite were attracted only by the specific odour of F. hispida . These results suggest that the use by non-pollinating fig wasps of a specific chemical message produced by figs could limit host shifts by non-pollinating fig wasps.     

Volatile cues used by the parasitoid, Aphytis melinus, for host location: California red scale revisited

Aphytis melinus DeBach (Hymenoptera: Aphelinidae) is a biological control agent of diaspidid scale insects. The parasitoid has a narrow host range but its hosts are polyphagous. We determined the source of volatile cues the wasp uses to locate its few host species when those hosts occur on more than one host plant species. We addressed four questions in regard to the use of volatile cues in host location of California red scale, Aonidiella aurantii (Maskell) (Homoptera: Diaspididae): (1) Does A. melinus use volatile cues to assist in host location? (2) Are these cues innately recognized or learned? (3) Are cues produced by female California red scale, or from other sources? (4) Are the cues specific to the host or host plant? These questions were tested through the use of a Y-tube olfactometer. Female A. melinus used volatile cues to orient toward both infested and uninfested host plant material. Wasps learned these cues by associating odors from the host plant with host presence. They had no innate preferences for scale insect or host plant volatile stimuli. Contrary to previous studies, we found no evidence of orientation toward the female-produced sex pheromone of California red scale, nor to volatile cues from the attacked host stage. Wasps given experience with scale insects growing on lemon fruit subsequently oriented toward lemon and orange fruit and leaves. The scale species with which the wasp was given experience did not affect this preference. Wasps given experience with California red scale growing on squash did not orient toward infested lemon fruit. The host ranges of the parasitoid and its hosts are used to explain the adaptive value for the evolution of learned rather than fixed responses to cues used in foraging behavior.     

Chemical mediation and niche partitioning in non-pollinating fig-wasp communities

1. The parasitic chalcidoid wasps associated with the species-specific and obligatory pollination mutualisms between Ficus spp. and their agaonid wasp pollinators provide a good model to study the functional organization of communities. 2. However, communities of non-pollinating fig wasps (NPFWs) remain little characterized, and their functioning and evolutionary dynamics are still poorly understood. 3. We studied the communities of NPFWs associated with the monoecious F. racemosa and the dioecious F. hispida. Associated with these two fig species are a total of seven wasp species belonging to three genera. These species present contrasts in life history traits and in timing of oviposition. The species studied are thus broadly representative of the communities of NPFWs associated specifically with fig-pollinator mutualisms. 4. In our study systems, there is temporal segregation of oviposition time among members of NPFW communities. 5. We tested the role of volatile chemicals in the attraction of NPFWs associated with these two fig species, and tried to determine if chemical mediation can explain the organization of the communities. 6. We conducted odour choice tests using a Y-tube olfactometer. All the NPFWs studied were shown to use volatile chemicals produced by the fig to locate their host. Furthermore, the signals used by each species depended on the phenological stage of the fig they exploit. 7. Results demonstrated that the pattern of oviposition results from the utilization of volatile signals produced by figs that vary in their composition at different stages of fig development. Thus, chemical mediation allows resource partitioning in the NPFW communities associated with fig-pollinator mutualisms, and suggests hypotheses to explain coexistence in other parasite communities.     

竹横锥大象对寄主及虫体挥发物的行为和触角电位反应

为了探明竹横锥大象Cyrtotrachelus buqueti Guerin-Meneville是否存在两性间的引诱作用,以及寄主植物慈竹Neosinocalamus affinis笋对两性行为的影响,采用Y型嗅觉仪测定了竹横锥大象雌、雄成虫对4种虫体及寄主挥发物不同处理（雌、雄成虫,慈竹笋与雌、雄成虫的复合体）的行为反应,测定了触角不同部位对雌雄成虫3种虫体的提取物（整体、鞘翅、后肠）及其与寄主植物挥发物质联合作用的触角电位反应（EAG）。结果表明：雌成虫挥发物对雄虫有较高的引诱活性,对雌虫有驱避作用;雄成虫挥发物对雌虫有一定的引诱作用。雌、雄成虫对虫体与竹笋复合体气味的行为和电生理反应均显著或极显著高于对相应虫体挥发物的反应。经EAG测定,触角各部位对虫体不同部位挥发物的EAG值差异明显,触角端部对虫体与植物挥发性物质的联合作用EAG反应差异极其显著（P＜0.01）。竹横锥大象对沾染雌雄成虫虫体提取物的玻璃棒有一定的兴奋表现,雌虫兴奋率为6.67%,雄虫为26.67%。结果提示,竹横锥大象种内存在信息素,雌虫释放的信息素对同类雄虫有很强的引诱作用,而雄虫所释放的信息素对同类雌虫有引诱作用,添加寄主植物能够增强两性间的引诱效果。     

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.