Contrasting performances of generalist and specialist Myzus persicae (Hemiptera: Aphididae) reveal differential prevalence of maternal effects after host transfer

Transgenerational maternal effects on performance (rm) after host transfer were evaluated in the generalist aphid Myzus persicae s.s., and in its subspecies specialized on tobacco, M. persicae nicotianae Blackman. We tested whether the performance of these taxa, when reared separately on optimal and suboptimal hosts (as sources of different maternal background) and then transferred to optimal hosts, experienced variations along four successive generations. Additionally, to compare the tolerance of both taxa to stress following host transfers, developmental instability (fluctuating asymmetry and body abnormalities) along the four generations was assessed. Taxon, rearing host, and generation affected the performance after host transfer. In the generalist, there was a significant improvement of rm along generations when transferred from suboptimal to optimal host and a significant decrease when transferred from optimal to optimal host; in the specialist, no increase or decrease occurred in any host transfer treatment. Transfer from suboptimal to optimal hosts caused higher losses of remaining replicates along generations than transfers from optimal to optimal hosts, and the specialist showed higher losses than the generalist. The only significant effect detected in comparisons involving fluctuating asymmetry values was that of taxon on length of siphunculi. Frequency of body abnormalities was not affected by treatments. Collectively, these results show a transgenerational weakening of maternal effects in the generalist but not in the specialist aphid, and suggest that rearing the latter in a suboptimal host causes not easily reversible changes that further give rise to constraints in performance.     

The tri-trophic interactions hypothesis: interactive effects of host plant quality, diet breadth and natural enemies on herbivores

Several influential hypotheses in plant-herbivore and herbivore-predator interactions consider the interactive effects of plant quality, herbivore diet breadth, and predation on herbivore performance. Yet individually and collectively, these hypotheses fail to address the simultaneous influence of all three factors. Here we review existing hypotheses, and propose the tri-trophic interactions (TTI) hypothesis to consolidate and integrate their predictions. The TTI hypothesis predicts that dietary specialist herbivores (as compared to generalists) should escape predators and be competitively dominant due to faster growth rates, and that such differences should be greater on low quality (as compared to high quality) host plants. To provide a preliminary test of these predictions, we conducted an empirical study comparing the effects of plant (Baccharis salicifolia) quality and predators between a specialist (Uroleucon macolai) and a generalist (Aphis gossypii) aphid herbivore. Consistent with predictions, these three factors interactively determine herbivore performance in ways not addressed by existing hypotheses. Compared to the specialist, the generalist was less fecund, competitively inferior, and more sensitive to low plant quality. Correspondingly, predator effects were contingent upon plant quality only for the generalist. Contrary to predictions, predator effects were weaker for the generalist and on low-quality plants, likely due to density-dependent benefits provided to the generalist by mutualist ants. Because the TTI hypothesis predicts the superior performance of specialists, mutualist ants may be critical to A. gossypii persistence under competition from U. macolai. In summary, the integrative nature of the TTI hypothesis offers novel insight into the determinants of plant-herbivore and herbivore-predator interactions and the coexistence of specialist and generalist herbivores.     

Host specialization in habitat specialists and generalists

Generalists and specialists use different cues to find their habitat and essential resources. While generalists have the advantage of exploiting a wider range of resources, they are predicted to be less efficient in using one particular resource compared to specialists. The level of specialization of parasitoids can be either at the habitat or at the host level; strategies used by either type are expected to differ. We examined interactions between three aphid parasitoid species that are a habitat specialist Aphidius rhopalosiphi, a habitat generalist Aphidius ervi, and a host generalist Praon volucre on three cereal aphids, Sitobion avenae, Metopolophium dirhodum and Rhopalosiphum padi. We compared total parasitism rate across behavioral and physiological variation in a non-choice test. Next, we addressed total parasitism in two phases to examine: (1) the response of parasitoids to different hosts through the behavioral sequence from antennation through oviposition, and (2) the physiological suitability of different hosts for oviposition and larval development. Parasitization typically involved the following behavioral steps: (1) antennal contact, (2) abdominal bending, and (3) ovipositor insertion (acceptance). A. rhopalosiphi had the same number of antennal contacts with the three aphids but showed fewer instances of abdominal bending towards R. padi. Pre-contact host preference was found for A. ervi but it did not correspond to the level of acceptance. The number of antennal contacts by P. volucre corresponded to the parasitization level of the aphid species but more mummies were produced on M. dirhodum than on R. padi. These results suggest that parasitoid species that are habitat specialists react similarly to the different host species present in the same habitat, whereas generalist species exhibit clear preferences during host selection. Preferences were, however, not always related to host suitability.     

Specialists make faster decisions than generalists: experiments with aphids

Theoretical studies and a few recent experimental reports suggest that the evolution of diet breadth in herbivorous insects is constrained by a limited neural ability to efficiently process large amounts of information in short periods of time. This neural-constraints hypothesis predicts that generalist herbivores should make slower or poorer decisions than specialists when selecting plants, because generalists must discriminate and decide among stimuli from a wider variety of potential hosts. The present study compares the speed with which host-associated decisions are made in specialist versus generalist populations of the aphid Uroleucon ambrosiae. Populations of U. ambrosiae from eastern North America are highly specific to the host plant Ambrosia trifida (Asteraceae), whereas those from the American southwest also feed on a variety of other taxa from the Asteraceae. Experiments with winged (alate) and wingless (apterous) individuals showed that host-finding, host-selection, host-acceptance, host-sampling and host-settling were more efficiently performed by the eastern specialists. These very consistent results provide evidence that strongly supports the neural-constraints hypothesis.     

The effect of rearing history and aphid density on volatile‐mediated foraging behaviour of Diaeretiella rapae

1. Parasitoid females foraging for hosts rely on cues derived from the insect host, the host plant and/or their interaction, and all of these can be learned during the immature and adult stages. 2. The present study investigated the importance of rearing history on foraging behaviour of Diaeretiella rapae, an endoparasitoid often associated with aphids feeding on brassicaceous plant species. 3. Parasitoids were reared on one of the four possible combinations, comprising two brassicaceous host plant species, Brassica nigra or Raphanus sativus, and two aphid species Brevicoryne brassicae or Myzus persicae. These parasitoids were tested in a Y‐tube olfactometer and given the choice between volatiles emitted by an aphid‐infested plant (25 or 100 aphids per plant) and an uninfested control plant. The parasitoid's responses were compared when offered: (i) the same plant–aphid combination as the one on which it had been reared; (ii) the same host plant infested with the alternative aphid species; or (iii) an alternative plant with the alternative aphid species. 4. Aphid density did affect the behavioural responses to the various odour sources, but rearing history did not. Diaeretiella rapae only preferred aphid‐induced to non‐induced plant volatiles at low aphid infestation level, whereas they did not discriminate between volatiles at high aphid infestation level. 5. It is concluded that aphid‐induced volatiles of brassicaceous plants play an important role during host habitat location, but seem less important for parasitoids to locate the aphid host itself. The data are discussed in the light of manipulation of host plant defences by aphids.     

Behavioural differences during host selection between alate virginoparae of generalist and tobacco-specialist Myzus persicae

Host plant selection and acceptance by aphids involves four consecutive steps: (1) prealighting behaviour, (2) leaf surface exploration and probing of subepidermal tissues, (3) deep probing of plant tissues, and (4) evaluation of the phloem sap. Host specialisation in aphids may involve not only different performances on potential hosts, but also different strategies for host selection and acceptance. Myzus persicae s.s. (Sulzer) (Homoptera: Aphididae) is one of the most polyphagous aphid species, although a tobacco‐adapted subspecies, M. persicae nicotianae, has been described. These two taxa constitute a good system for studying the effect of host range on host selection strategies. We studied the first two steps in the host selection process by alate virginoparae of M. persicae s.s. and M. persicae nicotianae on host and non‐host plants, using three types of behavioural assays: wind tunnel, olfactometry, and video‐recording. Alate virginoparae of M. persicae nicotianae recognised and chose their host plant more efficiently than M. persicae s.s., on the basis of olfactory and visual cues, and factors residing at cuticular and subcuticular levels. Host recognition was evident before phloem tissues were contacted. Olfactory cues were apparently not involved in host selection by M. persicae s.s.     

Evidence for predominant clones in a cyclically parthenogenetic organism provided by combined demographic and genetic analyses

Aphids are particularly interesting models in the study of genetic and demographic components of plant adaptation because of their breeding system which combines parthenogenesis and sexual reproduction (i.e. cyclical parthenogenesis), and the frequent emergence of host-adapted races reported in this group. In this paper, patterns of host adaptation were assessed on local populations of the aphid Sitobion avenae by following their demographic and genetic structure in a maize field for two consecutive years. The existence of putative generalist (polyphagous) or specialized (host-adapted) genotypes was also investigated by comparing the genotypic distribution of this aphid on maize and other cultivated host plants, using five microsatellite loci. Although population dynamics revealed strong variation in aphid abundance during the colonization period on maize, two genotypes identified at seven additional microsatellite loci were predominant and exhibited stable frequencies over cropping season and between years. Based on present and earlier studies, these two prevalent genotypes were shown to survive on different host plants other than maize, to colonize large geographical zones and to persist parthenogenetically for several years. All these data strongly suggest that these two genotypes are asexual generalist clones that could have been favoured by agricultural practices encountered in western Europe. Besides these two clones, a continual replacement of rare genotypes was observed on maize in both years. Hypotheses involving selection via aphid-plant interactions and natural enemies were proposed for explaining the disappearance of these genotypes on maize.     

豆柄瘤蚜茧峰的嗅觉定向反应(英文)

豆柄瘤蚜茧蜂Lysiphlebusfabarum对能寄生于包括大豆蚜Aphisglycines和豆蚜A.craccivora的多种蚜虫中。本实验研究了豆柄瘤蚜茧蜂在风洞中对以下气味源的嗅觉走向反应：未受损伤的寄主食料植株（UD）、机械损伤的寄主食料植株（MD）、蚜害植株（AD ,带蚜虫;AD～,不带蚜虫）、以及致害蚜虫。发现以豆蚜为寄主的豆柄瘤好茧蜂对AD 和AD~的正走向率高于对UD和MD的正走向率,以大豆蚜为寄主的豆柄瘤蚜虽峰对AD 的趋性高于对UD和大豆蚜。同时,豆柄瘤蚜茧蜂对UD、MD和致害蚜虫的定向反应率没有显著差异。这说明植食性昆虫的取食活动能诱导被害植株提高自身对害虫天敌的吸引力。此外还发现大豆是否受蚜害并不直接影响大豆蚜进行寄主定位。     

OLFACTORY ORIENTATION OF THE PARASITOID WASP LYSIPHLEBUS FABARUM TO ITS HOST FOOD PLANTS1)

Abstract  Lysiphlebus fabarum (Hymenoptera: Aphidiidae) is a generalist parasitic wasp species which can oviposit in several aphid species, including the bean aphid, Aphis craccivora (Homoptera Aphidinae), and the soybean aphid, A. glycines . The response of female L. fabarum and one of its host aphid species, A. glycines , to odors of undamaged host food plant (UD), mechanically damaged host food plant (MD), aphid-damaged plant (AD+, with aphid on, and AD˜, aphid washed off), and host aphid alone were studied in a wind tunnel. The positive orientation frequency of L. fabarum , which emerged from Aphis craccivora , to AD+ and AD^˜ was significantly greater than to the UD, MD. Similarily, L. fabarum that emerged from A. glycines significantly flew more often to AD+ than to UD and aphid alone. And there was no significant difference among the positive response frequency of L. fabarurn to UD, MD, and aphid alone. All these suggested that herbivore feeding induced the attraction of the damaged plant to natural enemy of herbivore. It is also found that the condition of soybean seedling, being damaged or undamaged, did not have direct impact on host plant location of aphid.     

Does rearing an aphid parasitoid on one host affect its ability to parasitize another species?

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Diet breadth and its relationship with genetic diversity and differentiation: the case of southern beech aphids (Hemiptera: Aphididae)

Herbivorous insect species with narrow diet breadth are expected to be more prone to genetic differentiation than insect species with a wider diet breadth. However, a generalist can behave as a local specialist if a single host-plant species is locally available, while a specialist can eventually behave as a generalist if its preferred host is not available. These problems can be addressed by comparing closely related species differing in diet breadth with overlapping distributions of insect and host populations. In this work, diet breadth, genetic diversity and population differentiation of congeneric aphid species from southern beech forests in Chile were compared. While at the species level no major differences in genetic diversity were found, a general trend towards higher genetic diversity as diet breadth increased was apparent. The aphid species with wider diet breadth, Neuquenaphis edwardsi (Laing), showed the highest genetic diversity, while the specialist Neuquenaphis staryi Quednau & Remaudière showed the lowest. These differences were less distinct when the comparisons were made in the same locality and over the same host. Comparison of allopatric populations indicates that genetic differentiation was higher for the specialists, Neuquenaphis similis Hille Ris Lambers and N. staryi, than for the generalist N. edwardsi. Over the same host at different locations, genetic differentiation among populations of N. edwardsi was higher than among populations of N. similis. The results support the assumption that specialists should show more pronounced genetic structuring than generalists, although the geographical distribution of host plants may be playing an important role.     

Behavior of bird cherry-oat aphid and green peach aphid in relation to potato virus Y transmission

Potato virus Y is transmitted to potato in a nonpersistent manner by many aphid species, some of which do not colonize this crop. The behavior of bird cherry-oat aphid, Rhopalosiphum padi (L.) on potato, Solanum tuberosum L., a plant species that is not colonized by this aphid, was described and compared with that of the potato-colonizing green peach aphid, Myzuspersicae (Sulzer). A higher proportion of winged morph of R. padi than M. persicae left the plant, but aphids that stayed in contact with the plant took the same mean time to initiate the first probe and it lasted the same mean time compared with M. persicae. Electronic penetration graph technique was used to study the probing behavior of the aphids during Potato virus Y (family Potyviridae, genus Potyvirus, PVY) transmission tests. Transmission rate decreased from 29 to 8% when the acquisition time increased from 5 min of continuous probing to 1 h with M. persicae, but it remained low (2 and 1%) with R. padi. Most of the difference in transmission rate between acquisition time with M. persicae and between aphid species was related to the change in the time and behavior taking place between the last cell puncture of the acquisition phase to the first cell puncture of the inoculation phase. Results presented here clearly demonstrated the importance of host plant selection and probing behavior in the transmission of nonpersistent plant viruses. They also stress the need to consider the behavior of the aphid in the design of laboratory tests of virus vector efficacy.     

Decision making by generalist and specialist aphids with the same genotype

One idea to explain the high incidence of specialisation in phytophagous insects is that their nervous systems are simple and unable to efficiently recognise multiple host plants. Here this 'neural limitations' hypothesis is tested using a generalist (the summer winged virginopara) and specialist (the autumn gynopara) morph of a single clone of the black bean aphid, Aphis fabae, that are almost identical morphologically and have the same genotype. Electrical monitoring of stylet behaviour and non-invasive behavioural observations are used to compare initial plant recognition, first reproduction, 'first registered phloem contact' and 'phloem acceptance' of the generalist and specialist on their specific host plants (generalist--herbaceous hosts, Beta vulgaris, Papaver dubium, Rumex obtusifolius, Vicia faba; specialist--woody host, Euonymus europaeus) and on the same host (E. europaeus). Additional analyses of behaviour over the first few minutes of plant contact are carried out for three mutual non-host plants: Euonymus alatus, Prunus padus and Brassica pekinensis. Few examples are found where the host-selection behaviour of the specialist on its host plant is more efficient than the generalist across its hosts and there is little difference in the behaviour of the different forms on the same host. Indeed there are several instances where the generalist is behaviourally more efficient than the specialist. These include absolute time to first reproduction by the generalist on various herbaceous hosts and the same host, E. europaeus, absolute time to first registered phloem contact by the generalist on P. dubium, and incidence of phloem acceptance by the generalist on several of its herbaceous hosts. Rejection of non-hosts is carried out more efficiently by the specialist gynopara on P. padus and B. pekinensis but not on E. alatus. For generalist and specialist morphs of the same A. fabae genotype, therefore, broad host range does not appear to be associated with a reduced ability to make plant-use decisions.     

Prey‐mediated effects of glucosinolates on aphid predators

1. Plant resistance against herbivores can act directly (e.g. by producing toxins) and indirectly (e.g. by attracting natural enemies of herbivores). If plant secondary metabolites that cause direct resistance against herbivores, such as glucosinolates, negatively influence natural enemies, this may result in a conflict between direct and indirect plant resistance. 2. Our objectives were (i) to test herbivore‐mediated effects of glucosinolates on the performance of two generalist predators, the marmalade hoverfly (Episyrphus balteatus) and the common green lacewing (Chrysoperla carnea) and (ii) to test whether intraspecific plant variation affects predator performance. 3. Predators were fed either Brevicoryne brassicae, a glucosinolate‐sequestering specialist aphid that contains aphid‐specific myrosinases, or Myzus persicae, a non‐sequestering generalist aphid that excretes glucosinolates in the honeydew, reared on four different white cabbage cultivars. Predator performance and glucosinolate concentrations and profiles in B. brassicae and host‐plant phloem were measured, a novel approach as previous studies often measured glucosinolate concentrations only in total leaf material. 4. Interestingly, the specialist aphid B. brassicae selectively sequestered glucosinolates from its host plant. The performance of predators fed this aphid species was lower than when fed M. persicae. When fed B. brassicae reared on different cultivars, differences in predator performance matched differences in glucosinolate profiles among the aphids. 5. We show that not only the prey species, but also the plant cultivar can have an effect on the performance of predators. Our results suggest that in the tritrophic system tested, there might be a conflict between direct and indirect plant resistance.     

Comparison of feeding behaviour of two Brassica pests Brevicoryne brassicae and Myzus persicae on wild and cultivated brassica species

Feeding behaviour of the specialist Brassicae aphid, Brevicoryne brassicae (L) (cabbage aphid) and the generalist, Myzus persicae, (Sulzer) (peach potato aphid) was monitored electronically on the susceptible cauliflower, Brassica oleracea var. botrytis cv Newton Seale, and a range of 17 Brassica species, B. carinata, B. juncea, B. nigra, B. macrocarpa, and B. villosa var. drepanensis and cultivated brassica varieties, B. oleracea, B. campestris and B. napus. Aphids, monitored for 10 h on the underside of leaves, performed recognisable feeding behaviour on all brassica species. The main differences in feeding behaviour, between M. persicae and B. brassicae, on the susceptible cauliflower Newton Seale, were fewer probes, shorter times to initially reach the phloem but longer times to establish sustained phloem ingestion and the longer times spent, by M. persicae, in xylem ingestion.Feeding behaviour on the range of brassica species tested indicated that generalist and specialist aphids are influenced differently by the host plant. A longer time spent in xylem ingestion was again the major difference in the feeding behaviour of the two aphids. In addition, rejection of passive phloem ingestion, by M. persicae, was not related so closely to increased time spent in non probing activities, as for B. brassicae. This observation indicates that M. persicae does not generally accept or reject brassica species due to the presence of phagostimulants, such as glucosinolates at the leaf surface or along the stylet pathway, unless the concentration is very high. Differences in feeding strategies employed by generalist and specialist aphids on the same plants are discussed.     

Canopy insect herbivores in the Azorean Laurisilva forests: key host plant species in a highly generalist insect community

This article explores patterns of insect herbivore distribution in the canopy of the Laurisilva forests on seven islands in the Azores archipelago. To our knowledge, this is one of the first extensive study of this type in tree or shrub canopies of oceanic island ecosystems. One of the most frequently debated characteristics of such ecosystems is the likely prevalence of vague, ill‐defined niches due to taxonomic disharmony, which may have implications for insect‐plant interactions. For instance, an increase in ecological opportunities for generalist species is expected due to the lack of predator groups and reduced selection for chemical defence in host plants. The following two questions were addressed: 1) Are specialists species rare, and insect herbivore species randomly distributed among host plant species in the Azores? 2) Are the variances in insect herbivore species composition, frequency and richness explained by host plants or by regional island effects? We expect a proportional distribution of herbivore species between host plants, influenced by host frequency and distinct island effects; otherwise, deviation from expectation might suggest habitat preference for specific host tree crowns. Canopy beating tray samples were performed on seven islands, comprising 50 transects with 1 to 3 plant species each (10 replicates per species), giving 1320 samples from ten host species trees or shrubs in total. From a total of 129 insect herbivore species, a greater number of herbivore species was found on Juniperus brevifolia (s=65) and Erica azorica (s=53). However, the number of herbivore species per individual tree crown was higher for E. azorica than for any other host, on all islands, despite the fact that it was only the fourth more abundant plant. In addition, higher insect species richness and greater insect abundance were found on the trees of Santa Maria Island, the oldest in the archipelago. Insect species composition was strongly influenced by the presence of E. azorica, which was the only host plant with a characteristic fauna across the archipelago, whereas the fauna of other plant crowns was grouped by islands. The great insect occurrence on E. azorica reflects strong habitat fidelity, but only four species were clearly specialists. Our findings indicate a broadly generalist fauna. The simplicity of Azorean Laurisilva contributed to the understanding of insect‐plant mechanisms in canopy forest habitats.     

Preference and performance of generalist and specialist herbivores on chemically defended host plants

1. Both the physiological efficiency (PE) hypothesis and the preference–performance (PP) hypothesis address the complex interactions between herbivores and host plants, albeit from different perspectives. The PE hypothesis contends that specialists are better physiologically adapted to their host plants than generalists. The PP hypothesis predicts that larvae perform best on the host plant preferred by ovipositing females. 2. This study tests components of both hypotheses using the specialist checkerspot, Euphydryas anicia, the generalist salt marsh caterpillar, Estigmene acrea, and host plants in the genus Penstemon, which are defended by iridoid glycosides. 3. In laboratory experiments, the generalist preferred and performed significantly better on the less well defended host plant species. This is consistent with results from a common garden experiment where the less well defended Penstemon species received more damage from the local community of generalists. Larvae of the specialist checkerspot preferred the more chemically defended species in the laboratory, but performed equally well on both hosts. However, field experiments demonstrated that adult checkerspot females preferred to oviposit on the less well defended host plant. 4. Components of the physiological efficiency hypothesis were supported in this system, as the specialist outperformed the generalist on the more iridoid glycoside‐rich host plant species. There was no support for the PP hypothesis, however, as there was no clear relationship between female preference in the field and offspring performance in the laboratory.     

Landscape structure and agricultural intensification are weak predictors of host range and parasitism rate of cereal aphids

Proportions of specialist and generalist primary parasitoids have been described by the resource breadth and the trade-off hypothesis. These alternative hypotheses predict either decreased or increased, respectively, parasitism rate of shared aphid species by specialist parasitoids. We tested both hypotheses and the confounding effects of landscape structure and agricultural intensification (AI) using extensive samplings of aphids and their parasitoids in Polish agricultural landscapes. Abundances, species composition of aphids, primary parasitoids, and parasitism rate of aphids by specialists and generalist parasitoids were analysed. Contrary to our expectations we found equally decreased parasitism rates by both types of primary parasitoids at higher aphid densities and thus proportion of specialists to generalists did not change with increasing host density. In line with the resource breadth hypothesis, specialist parasitoids had always lower abundances and parasitism rates than generalist parasitoids. Landscape diversity and agricultural intensification did not influence the host-parasitoid population dynamics. We speculate that these contrasting results could be caused by the additional density effects of secondary parasitoids. We conclude that simplistic two-trophic-level population models are not able to fully describe the complex dynamics of trophic networks. We also argue that agricultural intensification has lower effects on abundance and effectiveness of parasitoids than predicted by respective predator–prey models and empirical studies performed in controlled and artificial conditions.     

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.