Vector preference for hosts differing in infection status: sharpshooter movement and Xylella fastidiosa transmission

1. Epidemiological theory predicts that vector preference for hosts differing in infection status (i.e. healthy or infected) affects disease dynamics. 2. Numerous studies have documented strong vector preference for or discrimination against infected hosts. However, the significance of these behaviours for pathogen transmission and spread has been poorly described. 3. We conducted a series of choice assays to evaluate orientation preference, feeding preference, and movement rates of an important group of vectors, the sharpshooter leafhoppers, based on host infection status for the generalist plant pathogen, Xylella fastidiosa Wells et al. 4. Sharpshooters did not discriminate between healthy versus infected‐but‐asymptomatic grapevines, but they oriented preferentially to healthy grapevines more frequently than either symptomatic vines or those artificially coloured to mimic disease symptoms. 5. In a field trial three sharpshooter species showed different movement rates and preferences for feeding site, but all species exhibited similar and significant preference for healthy hosts. 6. Although there was no significant difference in acquisition efficiency among vector species, those individuals that spent more time on healthy hosts tended to be less likely to acquire the pathogen. 7. These results suggest that sharpshooters discriminate against infected grapevines, which are likely to be of poorer quality, with visual cues playing a role in host selection. Preference by these vectors may affect pathogen acquisition, which could affect disease spread in the field.     

Cucurbit aphid‐borne yellows virus (CABYV) modifies the alighting,settling and probing behaviour of its vector Aphis gossypii favouring its own spread

Plant pathogens are able to influence the behaviour and fitness of their vectors in such a way that changes in plant–pathogen–vector interactions can affect their transmission. Such influence can be direct or indirect, depending on whether it is mediated by the presence of the pathogen in the vector's body or by host changes as a consequence of pathogen infection. We report the effect that the persistently aphid‐transmitted Cucurbit aphid‐borne yellows virus (CABYV, Polerovirus) can induce on the alighting, settling and probing behaviour activities of its vector, the cotton aphid Aphis gossypii. Only minor direct changes on aphid feeding behaviour were observed when viruliferous aphids fed on non‐infected plants. However, the feeding behaviour of non‐viruliferous aphids was very different on CABYV‐infected than on non‐infected plants. Non‐viruliferous aphids spent longer time feeding from the phloem in CABYV‐infected plants compared to non‐infected plants, suggesting that CABYV indirectly manipulates aphid feeding behaviour through its shared host plant in order to favour viral acquisition. Viruliferous aphids showed a clear preference for non‐infected over CABYV‐infected plants at short and long time, while such behaviour was not observed for non‐viruliferous aphids. Overall, our results indicate that CABYV induces changes in its host plant that modifies aphid feeding behaviour in a way that virus acquisition from infected plants is enhanced. Once the aphids become viruliferous they prefer to settle on healthy plants, leading to optimise the transmission and spread of this phloem‐limited virus.     

Vector within‐host feeding preference mediates transmission of a heterogeneously distributed pathogen

1. Ecological theory predicts that vector preference for certain host species or discrimination between infected versus uninfected hosts impacts disease incidence. However, little information exists on the extent to which vector within‐host feeding preference mediates transmission. This may be particularly important for plant pathogens, such as sharpshooter transmission of the bacterium Xylella fastidiosa, which are distributed irregularly throughout hosts. 2. We documented the within‐host distribution of two vector species that differ in transmission efficiency, the leafhoppers Draeculacephala minerva and Graphocephala atropunctata, and which are free to move throughout entirely caged alfalfa plants. The more efficient vector D. minerva fed preferentially at the base of the plant near the soil surface, whereas the less efficient G. atropunctata preferred overwhelming the top of the plant. 3. Next we documented X. fastidiosa heterogeneity in mechanically inoculated plants. Infection rates were up to 50% higher and mean bacterial population densities were 100‐fold higher near the plant base than at the top or in the taproot. 4. Finally, we estimated transmission efficiency of the two leafhoppers when they were confined at either the base or top of inoculated alfalfa plants. Both vectors were inefficient when confined at the top of infected plants and were 20–60% more efficient when confined at the plant base. 5. These results show that vector transmission efficiency is determined by the interaction between leafhopper within‐plant feeding behaviour and pathogen within‐plant distribution. Fine‐scale vector and pathogen overlap is likely to be a requirement generally for efficient transmission of vector‐borne pathogens.     

Settling preferences of the whitefly vector Bemisia tabaci on infected plants varies with virus family and transmission mode

Virus infection may change not only the host‐plant phenotypic (morphological and physiological) characteristics, but can also modify the behavior of their insect vector in a mutualistic or rather antagonistic manner, to promote their spread to new hosts. Viruses differ in their modes of transmission and depend on vector behavior for successful spread. Here, we investigated the effects of the semi‐persistently transmitted Tomato chlorosis virus (ToCV, Crinivirus) and the persistent circulative Tomato severe rugose virus (ToSRV, Begomovirus) on alighting preferences and arrestment behavior of their whitefly vector Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) Middle East Asia Minor 1 (MEAM1) on tomato plants (Solanum lycopersicum L. cv. Santa Clara, Solanaceae). The vector alighting preferences between infected and uninfected plants in choice assays were apparently influenced by the presence of ToCV and ToSRV in the whiteflies or by their previous exposure to infected plants. The observed changes in vector behavior do not seem to benefit the spread of ToCV: non‐viruliferous insects clearly preferred mock‐inoculated plants, whereas ToCV‐viruliferous insects landed on mock‐inoculated and ToCV‐infected plants, indicating a partial change in insect behavior – ToCV was able to directly affect the preference of its vector B. tabaci, but this change in insect behavior did not affect the virus spread because viruliferous insects landed on mock‐inoculated and infected plants indistinctly. In contrast, ToSRV‐viruliferous insects preferred to land on mock‐inoculated plants, a behavior that increases the probability of spread to new host plants. In the arresting behavior assay, the majority of the insects remained on mock‐inoculated plants when released on them. A greater number of insects moved toward mock‐inoculated plants when initially released on ToCV‐ or ToSRV‐infected plants, suggesting that these viruses may repel or reduce the nutritional quality of the host plants for B. tabaci MEAM1.     

Induced Release of a Plant-Defense Volatile ‘Deceptively’ Attracts Insect Vectors to Plants Infected with a Bacterial Pathogen

Transmission of plant pathogens by insect vectors is a complex biological process involving interactions between the plant, insect, and pathogen. Pathogen-induced plant responses can include changes in volatile and nonvolatile secondary metabolites as well as major plant nutrients. Experiments were conducted to understand how a plant pathogenic bacterium, Candidatus Liberibacter asiaticus (Las), affects host preference behavior of its psyllid (Diaphorina citri Kuwayama) vector. D. citri were attracted to volatiles from pathogen-infected plants more than to those from non-infected counterparts. Las-infected plants were more attractive to D. citri adults than non-infected plants initially; however after feeding, psyllids subsequently dispersed to non-infected rather than infected plants as their preferred settling point. Experiments with Las-infected and non-infected plants under complete darkness yielded similar results to those recorded under light. The behavior of psyllids in response to infected versus non-infected plants was not influenced by whether or not they were carriers of the pathogen. Quantification of volatile release from non-infected and infected plants supported the hypothesis that odorants mediate psyllid preference. Significantly more methyl salicylate, yet less methyl anthranilate and D-limonene, was released by infected than non-infected plants. Methyl salicylate was attractive to psyllids, while methyl anthranilate did not affect their behavior. Feeding on citrus by D. citri adults also induced release of methyl salicylate, suggesting that it may be a cue revealing location of conspecifics on host plants. Infected plants were characterized by lower levels of nitrogen, phosphorus, sulfur, zinc, and iron, as well as, higher levels of potassium and boron than non-infected plants. Collectively, our results suggest that host selection behavior of D. citri may be modified by bacterial infection of plants, which alters release of specific headspace volatiles and plant nutritional contents. Furthermore, we show in a laboratory setting that this apparent pathogen-mediated manipulation of vector behavior may facilitate pathogen spread.     

Multiple guests in a single host: interactions across symbiotic and phytopathogenic bacteria in phloem‐feeding vectors – a review

Some pathogenic phloem‐limited bacteria are a major threat for worldwide agriculture due to the heavy economic losses caused to many high‐value crops. These disease agents – phytoplasmas, spiroplasmas, liberibacters, and Arsenophonus‐like bacteria – are transmitted from plant to plant by phloem‐feeding Hemiptera vectors. The associations established among pathogens and vectors result in a complex network of interactions involving also the whole microbial community harboured by the insect host. Interactions among bacteria may be beneficial, competitive, or detrimental for the involved microorganisms, and can dramatically affect the insect vector competence and consequently the spread of diseases. Interference is observed among pathogen strains competing to invade the same vector specimen, causing selective acquisition or transmission. Insect bacterial endosymbionts are another pivotal element of interactions between vectors and phytopathogens, because of their central role in insect life cycles. Some symbionts, either obligate or facultative, were shown to have antagonistic effects on the colonization by plant pathogens, by producing antimicrobial substances, by stimulating the production of antimicrobial substances by insects, or by competing for host infection. In other cases, the mutual exclusion between symbiont and pathogen suggests a possible detrimental influence on phytopathogens displayed by symbiotic bacteria; conversely, examples of microbes enhancing pathogen load are available as well. Whether and how bacterial exchanges occurring in vectors affect the relationship between insects, plants, and phytopathogens is still unresolved, leaving room for many open questions concerning the significance of particular traits of these multitrophic interactions. Such complex interplays may have a serious impact on pathogen spread and control, potentially driving new strategies for the containment of important diseases.     

Effect of host plant tissue on the vector transmission of grapevine leafroll-associated virus 3

Many biotic and abiotic factors affect the transmission efficiency of vector-borne plant pathogens. Insect vector within-plant distribution and host tissue preference are known to affect pathogen acquisition and inoculation rates. In this study, we first investigated whether feeding tissue affects the transmission of Grapevine leafroll-associated virus 3 by Planococcus ficus (Signoret) (Hemiptera: Pseudococcidae) and the effect of mealybug within-plant distribution on virus transmission under greenhouse conditions. Results showed no significant effect on transmission efficiency after insect confinement on leaf blades, petioles or stems of virus source or healthy test plants for either acquisition or inoculation trials. Transmission efficiency of a single mealybug varied from 4 to 25% in those trials. Second, we tested whether leaf position affected transmission efficiency due to potentially variable virus populations within acquisition plant tissues. No significant differences of transmission rate among acquisition leaf position were observed, probably because there were no differences in the virus population within source tissues. Finally, we examined the seasonality of the virus in field-collected samples and found that GLRaV-3 prevalence varied along a growing season, such that GLRaV-3 translocated along expanding shoots to leaves. Similarly, mealybug populations are known to increase in spring, and then mealybugs spread to cordons and leaves. This coordination of spatial and temporal dynamics of the virus and its vector may increase the risk of GLRaV-3 transmission during late spring and early summer. Further integration of information about pathogen populations in plants, vector feeding behavior and vector population seasonality could lead to more effective management practices.     

A powdery mildew infection on a shared host plant affects the dynamics of the Glanville fritillary butterfly populations

While biotrophic fungal pathogens have generally been considered to have negative effects on phytophagous insects sharing the same host plant, very little is known about whether fungal infection may affect the dynamics of natural insect populations. This study was designed to determine the effects of fungal infection by Podosphaera plantaginis , a powdery mildew, of a shared host plant, Plantago lanceolata , on the larvae of the butterfly Melitaea cinxia . Larval responses were assessed in a no-choice feeding assay involving infected and healthy leaves, as well as in a behavioural experiment in which larvae had an opportunity to move among infected and uninfected plants. In the no-choice feeding assay larvae developed more slowly and weighed less at diapause when feeding on fungal-infected than on healthy leaves. In the behavioural experiment larval groups tended to leave the original host plant when it was infected by P. plantaginis . This tendency was associated with splitting of larval groups into smaller subgroups. These effects observed in an experimental setting were also confirmed to act under natural conditions. An analysis of 167 M. cinxia populations showed that over-winter survival of larval groups was 26% lower in host populations infected by the mildew than in non-infected host populations. Smaller, more slowly developing larvae may not be ready to enter diapause at the onset of fall, causing the observed increase in mortality. This is the first study to demonstrate that the negative effects of a biotrophic fungal infection may extend to the dynamics of entire insect populations.     

Raspberry viruses affect the behavior and performance of Amphorophora agathonica in single and mixed infections

Pathogens may alter their hosts, which consequently increases transmission efficiency by vectors. We examined the effects of Raspberry leaf mottle virus [RLMV; Closterovirus (Closteroviridae)] and Raspberry latent virus [RpLV; Reovirus (Reoviridae)], alone and in a co‐infection in raspberry, Rubus idaeus L. (Rosaceae) cv. Meeker, on the behavior and performance of its vector, Amphorophora agathonica Hottes (Hemiptera: Aphididae). Longevity was increased in aphids feeding on all infected‐plant treatments compared with healthy plants, but aphid fecundity only increased in the co‐infection treatment. In a two‐way choice study between infected and healthy plants, aphids showed no difference in preference between plants after 30 min of exposure. After 24 h, aphids significantly preferred to settle on plants infected with RLMV over healthy, but healthy plants over plants infected with RpLV. There were no differences in settling preferences between healthy and co‐infected plants. An electrical penetration graph study showed no differences in aphid feeding behavior on plants infected with RLMV and RLMV+RpLV when compared with healthy controls. Our results are consistent with past findings that infected plant's impact vector performance and behavior, but also highlight the need to further investigate greater virus diversity and effects of mixed infections.     

Pathogens manipulate the preference of vectors,slowing disease spread in a multi‐host system

The spread of vector‐borne pathogens depends on a complex set of interactions among pathogen, vector, and host. In single‐host systems, pathogens can induce changes in vector preferences for infected vs. healthy hosts. Yet it is unclear if pathogens also induce changes in vector preference among host species, and how changes in vector behaviour alter the ecological dynamics of disease spread. Here, we couple multi‐host preference experiments with a novel model of vector preference general to both single and multi‐host communities. We show that viruliferous aphids exhibit strong preferences for healthy and long‐lived hosts. Coupling experimental results with modelling to account for preference leads to a strong decrease in overall pathogen spread through multi‐host communities due to non‐random sorting of viruliferous vectors between preferred and non‐preferred host species. Our results demonstrate the importance of the interplay between vector behaviour and host diversity as a key mechanism in the spread of vectored‐diseases.     

Plant virus infection modifies plant pigment and manipulates the host preference behavior of an insect vector

Insect-borne plant viruses may modify the phenotype of their host plants and thus influence the responses of insect vectors. When a plant virus modifies host preference behavior of a vector, it can be expected to influence the rate of virus transmission. In this study, we examined the effect of Maize Iranian mosaic virus (MIMV) infection on host preference behavior of the nymphs and adults of its vector, the small brown planthopper, Laodelphax striatellus Fallén (Hemiptera: Delphacidae), feeding on barley plants (Hordeum vulgare L., Poaceae). We found that both viruliferous nymphs and adults significantly preferred healthy plants, whereas non-viruliferous planthoppers preferred virus-infected barley. Further investigations revealed significant reductions in the chlorophyll and carotenoid contents of infected barley leaves. Based on these results, a possible association between insect host preferences and the pigment contents of the plants was observed. In summary, we suggest that host preference of L. striatellus could be affected by the propagative plant virus, possibly through association of this modification with some phenotypic traits of infected plants. These effects may have a critical impact on MIMV transmission rate, with significant implications for the development of virus epidemics.     

Influence of a propagative plant virus on the fitness and wing dimorphism of infected and exposed insect vectors

To maximize fitness, plant pathogenic viruses may manipulate their arthropod vectors through direct and indirect (via the host plant) interactions. For many virus-vector-plant associations, insect feeding does not always lead to virus acquisition. In fact, many plant viruses, especially those that propagate into their vectors, are acquired at low rates. Although the majority of insects colonizing an infected plant escape from viral infection, they are still exposed to the indirect effects (i.e. the effect of plant metabolism modification following virus infection). Little information has been reported on the effects of plant viruses on insects that become infected versus those that do not (here referred to as “exposed”). The effect that the Maize mosaic virus (MMV) (Rhabdoviridae) exerts on the fitness and wing dimorphism of the planthopper vector, Peregrinus maidis (Hemiptera, Delphacidae), that developed on leaves from either young or old corn plants was examined. MMV exerted non-consistent to minimal direct effects on developmental time, longevity, nymphal mortality and fecundity. In addition, some small yet significant fitness costs were encountered by exposed planthoppers to escape MMV infection. Furthermore, a significantly higher proportion of macropters over brachypters were produced on MMV-infected old leaves compared with healthy leaves of a similar age. We conclude that the virus influences the dispersal of the vector, promoting a larger production of macropters at the costs of brachypters at a late stage of the plant infection. Because MMV infection in planthoppers did not segregate by wing morphotype, our results indicate that the dispersal of both infected and exposed planthoppers was a likely consequence of the indirect effects of MMV.     

Influence of Xylella fastidiosa infection of citrus on host selection by leafhopper vectors

Infection of plants by pathogens can influence their attractiveness and suitability to insect vectors and other herbivores. Here we examined the effects of Citrus sinensis (L.) Osbeck (Rutaceae) infection by the bacterium Xylella fastidiosa, which causes citrus variegated chlorosis (CVC), on the feeding preferences of two sharpshooter vectors, Dilobopterus costalimai Young and Oncometopia facialis (Signoret) (Homoptera: Cicadellidae). Experiments were performed inside observation chambers, in which a healthy plant and an infected one (with or without CVC symptoms) were supplied to a group of 40 sharpshooters. The number of insects that selected each treatment was recorded at several time intervals in 48 h. In another experiment, the ingestion rate on healthy and infected (symptomatic or not) plants was evaluated by measuring the liquid excretion of sharpshooters that were confined on branches of each plant for 72 h. Both sharpshooter species preferred healthy plants to those with CVC symptoms. However, O. facialis did not discriminate between healthy citrus and symptomless infected plants. Feeding by D. costalimai was markedly reduced when confined on CVC‐symptomatic plants, but not on asymptomatic infected ones. The ingestion rate by O. facialis was not affected by the presence of CVC symptoms. The results suggest that citrus trees with early (asymptomatic) infections by X. fastidiosa may be more effective as inoculum sources for CVC spread by insect vectors than those with advanced symptoms.     

番茄褪绿病毒对MED烟粉虱成虫在不同寄主植物上的寄主选择性和取食行为的影响

【目的】由粉虱传播的番茄褪绿病毒(tomato chlorosis virus, ToCV)具有较强的暴发性和流行性,目前该病毒现已蔓延至世界各地,可为害多种作物,对农林经济生产造成严重危害。病毒侵染会影响介体昆虫的寄主选择性和取食行为,从而影响病毒传播。本研究旨在明确ToCV对MED烟粉虱Bemisia tabaci在不同寄主上的寄主选择性及取食行为的影响。【方法】利用Y型嗅觉仪测定携带和未携带ToCV的MED烟粉虱雌成虫的寄主选择性,利用刺吸电位(electrical penetration graph, EPG)技术比较携带和未携带ToCV的烟粉虱雌成虫在健康的番茄、辣椒、棉花和豇豆4种寄主植株上的取食行为差异。【结果】未携带ToCV的MED型烟粉虱雌成虫对于4种寄主植物的选择偏好性排序为番茄、辣椒>棉花>豇豆;携带ToCV烟粉虱雌成虫对4种寄主植物的选择性偏好性降低,排序为番茄、辣椒、棉花>豇豆。与未携带ToCV烟粉虱雌成虫相比,携带ToCV烟粉虱雌成虫取食4种植物所产生的刺探次数均显著增加,第1次到达韧皮部的时间有明显延后,取食总时间与韧皮部的取食时间明显减...     

Herbivore arthropods benefit from vectoring plant viruses

Plants infected with pathogens often attract the pathogens’ vectors, but it is not clear if this is advantageous to the vectors. We therefore quantified the direct and indirect (through the host plant) effects of a pathogen on its vector. A positive direct effect of the plant‐pathogenic Tomato spotted wilt virus on its thrips vector (Frankliniella occidentalis) was found, but the main effect was indirect; juvenile survival and developmental rate of thrips was lower on pepper plants that were damaged by virus‐free thrips than on unattacked plants, but such negative effects were absent on plants that were damaged and inoculated by infected thrips or were mechanically inoculated with the virus. Hence, potential vectors benefit from attacking plants with virus because virus‐infected plants are of higher quality for the vector's offspring. We propose that plant pathogens in general have evolved mechanisms to overcome plant defences against their vectors, thus promoting pathogen spread.     

Effect of Watermelon Silver Mottle Virus on the Life History and Feeding Preference of Thrips palmi

Thrips-borne tospoviruses cause numerous plant diseases that produce severe economic losses worldwide. In the disease system, thrips not only damage plants through feeding but also transmit causative agents of epidemics. In addition, thrips are infected with tospoviruses in the course of virus transmission. Most studies on the effect of tospoviruses on vector thrips have focused on the Tomato spotted wilt virus–Frankliniella occidentalis system. Thus, we focused on another thrips-borne tospovirus, Watermelon silver mottle virus (WSMoV), to examine the effect of virus infection on its vector, Thrips palmi. In this study, the direct and indirect effects of WSMoV on the life history traits and feeding preference of T. palmi were examined. The survival rate and developmental time of the WSMoV-infected larval thrips did not differ significantly from those of the virus-free thrips. Comparing the developmental time of larval thrips fed on the healthy plants, thrips-damaged plants, and thrips-inoculated plants (the WSMoV-infected plants caused by thrips feeding), feeding on the thrips-damaged plants reduced the developmental time, and the WSMoV infection in host plants partially canceled the effect of thrips damage on the developmental time. In addition, no significant variations between the virus-free and WSMoV-infected adult thrips regarding longevity and fecundity were observed. These results implied that WSMoV did not directly affect the life history traits of T. palmi, but the WSMoV infection indirectly affected the development of T. palmi through the virus-infected plants. Furthermore, feeding preference tests indicated that T. palmi preferred feeding on either the thrips-damaged plants or the thrips-inoculated plants to the healthy plants. The effect of tospoviruses on the life history and feeding preference of vector thrips might vary among host plants, virus species, vector species, and environmental factors.     

Prey infected with nucleopolyhedrovirus is preferred by the predatory stink bug Eocanthecona furcellata without altering its field fitness

Preference tests using the hemipteran predator Eocanthecona furcellata Wolff showed that nymphs and adults showed a constant preference towards virus-infected Spodoptera litura larvae. Adults feeding on healthy larvae throughout their life handled and consumed their prey quickly as compared to those feeding on infected prey, the differences being significant. Developmental time, survival rate, pre-mating period, sex-ratio and incubation period of E. furcellata that were reared on experimental plants infested with healthy larvae of S. litura as lifetime prey for all stages (control), healthy larvae during nymphal stage and infected larvae for adult stage, infected larvae for nymphal and healthy larvae for adult stage and infected larvae as lifetime meal for all stages of E. furcellata did not vary significantly. However, a significant reduction in body weight, fecundity, longevity and percent egg hatchability was observed when E. furcellata were fed on infected larvae as lifetime meal. The results revealed that the virus-infected larvae, relative to healthy larvae, had significant effect on prey preference of predators, but feeding on virus-infected prey had no adverse effect on its field fitness except on lifetime meal, which may not hold true under field conditions.     

Temporal Effects of a Begomovirus Infection and Host Plant Resistance on the Preference and Development of an Insect Vector,Bemisia tabaci,and Implications for Epidemics

Persistent plant viruses, by altering phenotypic and physiological traits of their hosts, could modulate the host preference and fitness of hemipteran vectors. A majority of such modulations increase vector preference for virus-infected plants and improve vector fitness, ultimately favouring virus spread. Nevertheless, it remains unclear how these virus-induced modulations on vectors vary temporally, and whether host resistance to the pathogen influences such effects. This study addressed the two questions using a Begomovirus-whitefly-tomato model pathosystem. Tomato yellow leaf curl virus (TYLCV) -susceptible and TYLCV-resistant tomato genotypes were evaluated by whitefly-mediated transmission assays. Quantitative PCR revealed that virus accumulation decreased after an initial spike in all genotypes. TYLCV accumulation was less in resistant than in susceptible genotypes at 3, 6, and 12 weeks post inoculation (WPI). TYLCV acquisition by whiteflies over time from resistant and susceptible genotypes was also consistent with virus accumulation in the host plant. Furthermore, preference assays indicated that non-viruliferous whiteflies preferred virus-infected plants, whereas viruliferous whiteflies preferred non-infected plants. However, this effect was prominent only with the susceptible genotype at 6 WPI. The development of whiteflies on non-infected susceptible and resistant genotypes was not significantly different. However, developmental time was reduced when a susceptible genotype was infected with TYLCV. Together, these results suggest that vector preference and development could be affected by the timing of infection and by host resistance. These effects could play a crucial role in TYLCV epidemics.     

Multitrophic interactions of the silverleaf whitefly,host plants,competing herbivores,and phytopathogens

Our laboratory found that silverleaf whitefly (SLW; Bemisia argentifolii Bellows & Perring) feeding alters host plant physiology and chemistry. The SLW induces a number of host plant defenses, including pathogenesis-related (PR) protein accumulation (e.g., chitinases, beta-1,3-glucanases, peroxidases, chitosanases, etc.). Induction of the PR proteins by SLW feeding occurs in various plant species and varieties. The extent and type of induction is dependent on a number of factors that include host plant growing conditions, the length of time the host plant is exposed to SLW feeding, the plant variety, and SLW population densities. The appearance of PR proteins correlates well with reduced infestations of conspecific insect herbivore competitors. Greenhouse and field experiments in which herbivore competitors (cabbage looper, Trichoplusia ni; leaf miner, Liromyza trifolii) were placed on plants previously exposed to SLW feeding demonstrated behavioral differences (oviposition, feeding preferences) and reduced survival rates and development times of these insects. The interaction was asymmetrical, i.e., SLW infestations of plants previously exposed to leaf miners had little or no effect on SLW behavior (oviposition). Induction of plant-defensive proteins by SLW feeding was both local (at the feeding site) and systemic (uninfested leaves distant to the feeding site). There are interactions between diseases such as tomato mottle virus (ToMoV; a geminivirus) and the host plant and SLW. PR proteins were induced in tomato plants infected with ToMoV much as they were via non-viruliferous SLW feeding. The presence of ToMoV in tomato plants significantly increased the number of eggs produced by SLW females. Experiments using tomato plants, powdery mildew (PM), and tobacco mosaic virus (TMV) show that whitefly infestations can affect plant pathogen relationships but the effects vary among pathogen types. Enzyme analyses prior to pathogen inoculation showed that whitefly treatment significantly increased the activities of foliar chitinase and peroxidase. Evaluation of pathogen growth 3 weeks after inoculation showed that whitefly feeding significantly reduced the incidence of PM. However, TMV levels evaluated by ELISA were not significantly affected by whitefly feeding. Six weeks after inoculation with pathogens, the chitinase and peroxidase activities were still elevated in plants initially fed on by whiteflies but continuing pathogen infection had no effect on these enzymes. The possibility that geminivirus infection and/or SLW infestations isolate the host plant for the selected reproduction of the virus and the insect is discussed. Multitrophic cascade effects may contribute to the successful eruptive appearance of SLW on various crops, ranking them as a major pest. They may explain the general observation that when SLW infest a host plant there are few if any competing insect herbivores and pathogens found in the host. However, the results indicate that certain SLW-virus relationships could be mutualistic.     

Prior flea beetle herbivory affects oviposition preference and larval performance of a potato beetle on their shared host plant

Abstract 1. The herbaceous plant Solanum carolinense (L.) (Solanaceae) is host to a number of specialist insects, including the leaf-feeding beetles Epitrix fuscula (Crotch) and Leptinotarsa juncta (Germar) (Coleoptera: Chrysomelidae). Potted individuals of S. carolinense were subjected to one of two treatments: exposure to herbivory by E. fuscula or exclusion of all herbivores. The effects of E. fuscula herbivory on larval performance and oviposition preference of L. juncta were investigated.
2. Although the masses of the L. juncta pupae did not differ between the two treatments, larvae feeding on damaged plants developed more slowly than those feeding on undamaged plants.
3. In both paired leaf choice trials and whole plant choice trials, larvae of L. juncta showed no preference for undamaged versus damaged hosts.
4. In a field transplant experiment, adult L. juncta females showed slight feeding preferences and strong oviposition preferences for undamaged plants versus plants that had been fed on by E. fuscula .
5. The results are discussed with reference to their implications for plant-mediated competition among herbivores and constraints on the evolution of plant resistance.