Leaf miner and plant galler species richness on Acacia: relative importance of plant traits and climate

Diversity patterns of herbivores have been related to climate, host plant traits, host plant distribution and evolutionary relationships individually. However, few studies have assessed the relative contributions of a range of variables to explain these diversity patterns across large geographical and host plant species gradients. Here we assess the relative influence that climate and host plant traits have on endophagous species (leaf miners and plant gallers) diversity across a suite of host species from a genus that is widely distributed and morphologically variable. Forty-six species of Acacia were sampled to encapsulate the diversity of species across four taxonomic sections and a range of habitats along a 950 km climatic gradient: from subtropical forest habitats to semi-arid habitats. Plant traits, climatic variables, leaf miner and plant galler diversity were all quantified on each plant species. In total, 97 leaf mining species and 84 plant galling species were recorded from all host plants. Factors that best explained leaf miner richness across the climatic gradient (using AIC model selection) included specific leaf area (SLA), foliage thickness and mean annual rainfall. The factor that best explained plant galler richness across the climatic gradient was C:N ratio. In terms of the influence of plant and climatic traits on species composition, leaf miner assemblages were best explained by SLA, foliage thickness, mean minimum temperature and mean annual rainfall, whilst plant gall assemblages were explained by C:N ratio, %P, foliage thickness, mean minimum temperature and mean annual rainfall. This work is the first to assess diversity and structure across a broad environmental gradient and a wide range of potential key climatic and plant trait determinants simultaneously. Such methods provide key insights into endophage diversity and provide a solid basis for assessing their responses to a changing climate.     

Plant extracts as an alternative to control Leucoptera coffeella (Guérin-Mèneville) (Lepidoptera: Lyonetiidae)

We evaluated the effects of crude extracts from the plantain Plantago lanceolata and the bitter gourd Momordica charantia on the oviposition preference and development of the coffee leaf miner Leucoptera coffeella Guérin-Mèneville & Perrottet under laboratory and/or greenhouse conditions. The ovicidal effects of these extracts were also studied in a greenhouse. Plantago lanceolata and M. charantia extracts also underwent fractionation directed by oviposition tests with the coffee leaf miner. The extracts of both plants reduced L. coffeella oviposition and egg hatching, apparently as a result of action of plant metabolites on the embryo. Adults originating from eggs treated with the extracts exhibited similar survival rates, but a higher female/male ratio. Fecundity was reduced for females obtained from eggs treated with the M. charantia extract. Partial chemical analysis indicated that both extracts produced polar fractions that reduced the oviposition of L. coffeella on coffee leaves under laboratory conditions. The extracts of P. lanceolata and M. charantia have potential for use in the development of new products to control the coffee leaf miner.     

Determining Plant – Leaf Miner – Parasitoid Interactions: A DNA Barcoding Approach

A major challenge in network ecology is to describe the full-range of species interactions in a community to create highly-resolved food-webs. We developed a molecular approach based on DNA full barcoding and mini-barcoding to describe difficult to observe plant – leaf miner – parasitoid interactions, consisting of animals commonly regarded as agricultural pests and their natural enemies. We tested the ability of universal primers to amplify the remaining DNA inside leaf miner mines after the emergence of the insect. We compared the results of a) morphological identification of adult specimens; b) identification based on the shape of the mines; c) the COI Mini-barcode (130 bp) and d) the COI full barcode (658 bp) fragments to accurately identify the leaf-miner species. We used the molecular approach to build and analyse a tri-partite ecological network of plant – leaf miner – parasitoid interactions. We were able to detect the DNA of leaf-mining insects within their feeding mines on a range of host plants using mini-barcoding primers: 6% for the leaves collected empty and 33% success after we observed the emergence of the leaf miner. We suggest that the low amplification success of leaf mines collected empty was mainly due to the time since the adult emerged and discuss methodological improvements. Nevertheless our approach provided new species-interaction data for the ecological network. We found that the 130 bp fragment is variable enough to identify all the species included in this study. Both COI fragments reveal that some leaf miner species could be composed of cryptic species. The network built using the molecular approach was more accurate in describing tri-partite interactions compared with traditional approaches based on morphological criteria.     

Modelling the distribution of Ilex aquifolium at the north-eastem edge of its geographical range

Plant distribution borders are key features to characterise the ecological niche of a species and to monitor effects of climate change. Here we focus on an evergreen small tree, Ilex aquifolium, which reaches its north-eastern range edge in Denmark. Our main objectives are to describe and to model the current distribution of the species, to identify the most important climatic and land use factors which shape this distribution pattern, and to analyse the species' habitat requirements. For this purpose we used data from a national mapping project, complemented by information from forest owners. The distribution and abundance of I. aquifolium in Denmark have markedly changed during the past 40 years. It is now found in almost all districts, although the centres of abundance still coincide with the historical records. Our model shows lower habitat suitability for the species in northern and eastern districts, where winters are more severe and spring precipitation is lower. To a lesser extent, land use influences I. aquifolium occurrence, but it is more common in areas with a high proportion of forests and/or urban sites. The analysis of habitat requirements supports these results, since I. aquifolium occurs mainly as a forest species in deciduous stands, on relatively nutrient-rich moist soils, and under moderately high light conditions. However, some records may be the product of seed dispersal from planted individuals nearby. The results suggest that the range edge of the species has moved at least 100 km east within half a century. Since I.aquifolium is sensitive to winter frost, this change in distribution may be due to increasingly mild winter temperatures.     

Volatiles that encode host-plant quality in the grapevine moth

Plant volatiles are signals used by herbivorous insects to locate host plants and select oviposition sites. Whether such volatiles are used as indicators of plant quality by adult insects in search of host plants has been rarely tested. We tested whether volatiles indicate plant quality by studying the oviposition of the grapevine moth Lobesia botrana on the grapevine plant Vitis vinifera. Host plants were infected with a variety of microorganisms, and larval fitness was correlated to the infected state of the substrate. Our results show an oviposition preference for volatiles that is significantly correlated with the fitness of the substrate. The chemical profiles of the bouquets from each V. vinifera–microorganism system are clearly differentiated in a PCA analysis. Both the volatile signal and the quality of the plant as larval food were affected by the introduction of microorganisms. Our study represents a broad approach to the study of plant–insect interactions by considering not only the direct effect of the plant but also the effect of plant–microorganism interactions on insect population dynamics.     

Climate change effects on physiology and population processes of hosts and vectors that influence the spread of hemipteran-borne plant viruses

Plant virus diseases constitute one of the limiting factors to the productivity of agriculture. Changes in host plants and insect vector populations that might result from climate change (their geographical distribution range, their densities, migration potential and phenology) could affect the spread of plant viruses. At the individual level, alterations in plant physiological processes that are relevant to their molecular interactions with viruses, like changes in metabolism, leaf temperature, and their effects on some processes, like the temperature-sensitive antiviral resistance based in RNA silencing, can also influence the ability of individual plants to control viral infections. In order to assess the impact that climate change may have on the incidence and spread of hemipteran-borne plant viruses, its potential effects on virus/plant interactions and hemipteran insect vectors, as well as other operating processes, which could exacerbate or mitigate them, are identified and analyzed in this review.     

Leaf trait variation on Erythroxylum tortuosum (Erythroxylaceae) and its relationship with oviposition preference and stress by a host‐specific leaf miner

It has been suggested that plant physical and chemical traits vary considerably in space and time. Hence, leaf‐mining insects may adjust their oviposition in response to leaf attributes representing high quality. Moreover, herbivorous insects can modify leaf morphology by acting as stressors, increasing, for example, fluctuating asymmetry (FA) levels. Here, we investigate oviposition preference in Agnippe sp.2, a leaf‐mining moth of Erythroxylum tortuosum, in relation to differences in leaf nutritional quality (i.e. levels of water, nitrogen and tannin content), leaf area (i.e. quantity of resource hypothesis) and FA. We also verify whether temporal variation in plant nutritional quality emerges as an alternative hypothesis to explain oviposition distribution in time, and whether this leaf miner is a stress‐causing agent, increasing FA during larval development. Mined leaves and leaves with and without eggs were periodically collected from plants located in a Cerrado fragment in Brazil. In the laboratory, leaf traits were assessed (using image analysis software) and quantified (biochemical analysis) according to the aims previously determined. Oviposition probability did not change in relation to variations in nitrogen, tannins and FA of leaves. However, leaf‐miner females preferred to oviposit on leaves having large areas and low water contents. It was also verified that new leaves of E. tortuosum, which carried most leaf‐miner eggs, presented significantly lower tannins and greater levels of nitrogen and water than old leaves. The oviposition choice exhibited by leaf miners was found to be non‐random because they appear to use resource quantity and water content as cues as where to lay their eggs. The temporal variation of plant nutritional quality is likely to influence the time of leaf‐miner oviposition; and leaf FA was not increased during larval feeding, suggesting that these herbivores do not cause variations in FA levels.     

Influence of plant genotype and environment on oviposition preference and offspring survival in a gallmaking herbivore

Summary Plant resistance to insect herbivores may derive from traits influencing herbivore preference, traits influencing the suitability of the plant as a host, or both. However, the plant traits influencing host-plant selection by ovipositing insect herbivores may not completely overlap those traits that affect larval survival, and distinct traits may exhibit different levels of genetic vs. environmental control. Therefore, resource supply to the host plant could affect oviposition preference and larval performance differently in different plant genotypes. To test this hypothesis, the effects of resistance level, plant genotype, and resource supply to the host plant on oviposition preference and larval performance of a gallmaking herbivore, and on various plant traits that could influence these, were examined. Replicates of four genotypes of Solidago altissima, grown under low, medium, or high levels of nutrient supply in full sun or with medium levels of nutrients in shade, were exposed to mass-released Eurosta solidaginis. The number of plants ovipunctured was significantly affected by plant genotype and the interaction between genotype and nutrient supply to the host plant: one susceptible and one resistant genotype were more preferred, and preference tended to increase with nutrient supply in the more-preferred genotypes. The growth rate of ovipunctured plants during the oviposition period was significantly greater than that of unpunctured plants. Bud diameter (which was strongly correlated with plant growth rate), leaf area, and leaf water content were significant determinants of the percentage of plants ovipunctured, explaining 74% of the variance. The number of surviving larvae was significantly affected by plant genotype, but no effect of nutrient or light supply to the host plant was detected. The ratio of bud diameter to bud length was positively related to the percentage of ovipunctured plants that formed galls, suggesting that the accurate placement of eggs near the apical meristem by ovipositing females may be easier in short, thick buds. No significant correlation was observed between oviposition preference and larval survival at the population level. These results suggest that the plant traits affecting oviposition preference may exhibit different magnitudes of phenotypic plasticity than those affecting larval survival, and that the degree of phenotypic plasticity in plant traits affecting oviposition preference may differ among genotypes within a species.     

Effect of arbuscular mycorrhizal fungi (Glomus intraradices) on the oviposition of rice water weevil (Lissorhoptrus oryzophilus)

Root-feeding insects are important drivers in ecosystems, and links between aboveground oviposition preference and belowground larval performance have been suggested. The root-colonizing arbuscular mycorrhizal fungi (AMF) play a central role in plant nutrition and are known to change host quality for root-feeding insects. However, it is not known if and how AMF affect the aboveground oviposition of insects whose offspring feed on roots. According to the preference–performance hypothesis, insect herbivores oviposit on plants that will maximize offspring performance. In a greenhouse experiment with rice (Oryza sativa), we investigated the effects of AMF (Glomus intraradices) on aboveground oviposition of rice water weevil (Lissorhoptrus oryzophilus), the larvae of which feed belowground on the roots. Oviposition (i.e., the numbers of eggs laid by weevil females in leaf sheaths) was enhanced when the plants were colonized by AMF. However, the leaf area consumed by adult weevils was not affected. Although AMF reduced plant biomass, it increased nitrogen (N) and phosphorus concentrations in leaves and N in roots. The results suggest that rice water weevil females are able to discriminate plants for oviposition depending on their mycorrhizal status. The discrimination is probably related to AMF-mediated changes in plant quality, i.e., the females choose to oviposit more on plants with higher nutrient concentrations to potentially optimize offspring performance. AMF-mediated change in plant host choice for chewing insect oviposition is a novel aspect of below- and aboveground interactions.     

Biology and Host Range of the Green-seed Weevil, Sibinia fastigiata , for Biological Control of Mimosa pigra

Aspects of the biology and host range of Sibinia fastigiata Clark (Coleoptera: Curculionidae) were studied to assess its safety for release in Australia as a biological control agent of the weed Mimosa pigra L . (Mimosaceae) . Larvae feed on the seeds and adults on open flowers of their host . Adults oviposit on to immature seeds 3 mm long or less and hence seeds of this length and maturity were used in the host range tests and for rearing . Females are shown to avoid previously attacked seeds enhancing their effectiveness as seed destroyers . Survival of adults was higher when provided with open flowers . The host range was determined using laboratory control - choice oviposition tests on excised plant material and , in the field in the native range , no - choice oviposition tests on living plants , surveys of adults on plants , and breeding of insects from pods of plants of various legume species . The control - choice oviposition tests employed a new design in which the control plant alone was offered to the insects followed by a choice of test plants species . Other than M. pigra, only one plant species was acceptable for oviposition , the closely related M. asperata. Larval development also occurs on M. asperata and this host is occasionally used in the field . This insect was approved for release in March 1997 .     

Physical barriers and corridors in urban habitats affect colonisation and parasitism rates of a specialist leaf miner

1. Urban environments are fragmented habitats characterised by the presence of physical barriers, which may negatively affect dispersal and colonisation by insect herbivores and their natural enemies. Conversely, plants growing along pavements may function as dispersal corridors, helping to moderate the harmful effects of resource patch isolation on organism movement and population persistence. 2. We experimentally tested the effects of walls as physical barriers to the dispersal of the leaf miner Liriomyza commelinae Frost and colonisation of its host plant, Commelina erecta L., in urban habitats. We also evaluated whether plants along pavements could act as corridors for this species. 3. We exposed experimental host plants to the leaf miner in houses with front gardens and back yards, the latter being completely surrounded by walls. The front gardens had walls but none separating them from the pavement. Previously mined plants were also exposed to parasitoids in the yards to determine parasitoid attack. 4. Liriomyza commelinae took longer to colonise back yards with higher walls, and the abundance of mined plants along pavements reduced the colonisation time. Leaf‐miner abundance was marginally affected by the yard type, and was lower in back yards. Cumulative parasitism rates decreased with increasing distance at which mined plants were placed from pavements. 5. Constructions act as physical barriers, having a negative impact on colonisation of host plants by leaf miners. The function of pavements as corridors seems to depend on the abundance of mined plants. Parasitism may be affected by distance from the corridor rather than physical barriers or other potential hosts.     

Suitability of stressed and vigorous plants to various insect herbivores

We conducted a controlled experiment to test the plant vigor and the plant stress hypotheses. The two hypotheses associate plant physiological conditions to insect feeding mode and performance. We exposed tomato, Lycopersicon esculentum, to different types of growing conditions: optimal (vigorous plants), resource based stress (water and/or nutrient deficit), and physical stress (punched hole in terminal leaflets). Plant performance, foliar nutritional value for insects and chemical defenses were analyzed after 14 d. These plants were offered to insects belonging to distinct feeding guilds: the silverleaf whitefly, Bemisia argentifolii, a phloem feeder; the leafminer, Liriomyza trifolii; and the corn earworm, Heliothis zea, a leaf chewing caterpillar.
The experimental conditions generated a gradient of plant growth in the following order: optimal (vigorous)>control=hole punched>no fertilizer>no water>no water and no fertilizer. The last two treatments resulted in plants with poor nutritional value (based on %water, C/N, %N) and higher levels of defensive compounds (i.e., peroxidase and total phenolics) compared with control and the vigorous plants. Hole‐punching neither affected plant growth nor any of the phytochemicals measured. In a choice experiment adult whitefly ovipositioning was not affected by either vigor or punching but was reduced on the other plants (P<0.01). Leafminer feeding and oviposition and corn earworm larval growth rates were higher on the vigorous plants and lower on the punched, no fertilizer, no water, and no water and no fertilizer host plants (P<0.01).
Regardless of insect species or bioassay method, the results in the tomato system support the plant vigor hypothesis that predicts positive association between insect performance and plant growth. The results contradict the plant stress hypothesis that rank stressed plants as better hosts for insects. The mechanisms involved are a combination of poor nutritional value and chemical defenses. We demonstrate a negative association between plant growth and chemical defense. However, induced response triggered by hole‐punching was not cost effective to the plants.     

Plant-mediated effects of drought stress on host preference and performance of a grass miner

The plant stress hypothesis predicts that environmental stress increases the suitability of plants as food for herbivores, especially for senescence feeders. Yet, performance is enhanced only at moderate stress intensities in several herbivores. Even more paradoxically, a large number of insect species prefer and perform better on vigorously growing plants. In order to test plant stress theory, we conducted a laboratory experiment in which the influence of plant water stress on host preference and the performance of the grass miner Chromatomyia milii was studied. We imposed a gradient of stress intensities, i.e. 25, 50, 75, 150, and 300‐ml weekly‐administered water per grass pot, in order to study the full range of responses of C. milii to water stressed plants. Plant stress intensity was quantified by measuring individual plant mass, foliar water content and the concentration of the photosynthetic pigments chlorophyll a and b. Plant mass had decreased from the 150 and 300‐ml treatments to the lowest water treatment at the end of our experiment, which was mainly a result of a reduction in leaf area and leaf number. Foliar water content was clearly negatively affected by water shortage. Chlorophyll a and b also decreased with water shortage. Finally, the stress intensity measurements showed that plants acclimated to water stress conditions throughout the experiment. Feeding and oviposition preference of C. milii was positively related to water supply. No larvae survived on two lowest water treatments and only 38% survived on the 75‐ml treatment, while more than 80% survived on the 150 and the 300‐ml treatments. Offspring development time was longer on the 75‐ml treatment than on the 150 and the 300‐ml treatments. We also evaluated the mechanisms that could explain the response of C. milii to water stressed plants. Although no relationship between water treatment and foliar amino acid concentration was found, we observed significantly higher foliar protein concentrations in the 25 and the 50‐ml treatments. This supports the hypothesis that abiotic stress causes an increase of nitrogenous compounds in plants. Leaf senescence following self‐pruning, a process by which H. lanatus plants acclimate to drought conditions, was responsible for the dramatically high offspring mortality on the water stressed plants. The shape of the plant stress intensity–herbivore response relationship showed strong variation and depended both on the type of plant stress intensity measure and herbivore response variable involved. Yet, all relationships showed a monotonic increase of herbivore preference and performance with decreasing plant stress intensity. This indicates that C. milii prefers and performs better on vigorously growing plants. We found no support for an increased herbivore performance on moderately or severely stressed plants.     

Risk to non-target plants from Charidotis auroguttata (Chrysomelidae: Coleoptera), a potential biocontrol agent for cat’s claw creeper Macfadyena unguis-cati (Bignoniaceae) in Australia

The potential of the leaf beetle Charidotis auroguttata as a biocontrol agent for cat’s claw creeper Macfadyena unguis-cati (Bignoniaceae), an environmental weed in Australia, and risk to non-target plants was evaluated under quarantine conditions. In no-choice tests, C. auroguttata adults and larvae fed on many plant species across different families, but egg to adult development occurred only on the target weed. However, when neonate larvae from the target weed were transferred onto Myoporum boninense australe (Myoporaceae), a non-target native plant, 11.7% completed development, as compared to 95% of larvae that completed development on the target weed. Larval development on this non-target species also took twice as long as on the target weed. No larvae completed development on other test plants. In choice tests, leaf area consumption by adults and larvae was significantly more on the target weed than on other plants, and oviposition occurred only on the target weed. In the no-choice demography trials, adults laid eggs from the second week after emergence on the target weed, with an average of 0.286 eggs/female/day, resulting in an 18-fold increase in the adult population over 16 weeks. On My. boninense australe adult survival remained high, but oviposition commenced only from the 10th week after emergence with an average of 0.023 eggs/female/day, and none of the eggs developed into adults. In the choice demography trials, oviposition on the target weed was evident from the fourth week onwards, while on the non-target plant oviposition commenced only from the 14th week. Only 10% of total adults and 11.3% of total eggs were found on the non-target plant, and none of these eggs developed into adults. Although the biocontrol agent can ‘spill-over’ from the target weed to the non-target native plant and cause adult feeding damage, the non-target plant could not sustain a viable insect population on its own. This agent was not approved for field release in Australia due to perceived risk to non-target species.     

Plant‐mediated interactions among above‐ground and below‐ground life stages of a root‐feeding weevil

1. Interactions among herbivores mediated by plant responses to herbivore injury may have large impacts on herbivore population densities. Responses may persist for weeks after injury and may affect not only the initial (inducing) herbivore, but also herbivores that are spatially or temporally separated from the initial attacker. 2. In many plant–insect interactions, multiple life stages of the insect may be associated with the same plant, and these various stages may interact indirectly with one another via induced responses. The rice water weevil (RWW), Lissorhoptrus oryzophilus, a serious global pest of rice, is one such insect. A series of experiments were performed with root‐feeding larvae and leaf‐feeding adults of the RWW using three conventional rice varieties. 3. The first objective of this study was to test whether RWW adult feeding on rice leaves resulted in altered oviposition by subsequent adults. The hypothesis for the first objective was that RWW adult feeding would decrease plant suitability, resulting in reduced oviposition by subsequent adults. 4. The second objective was to test whether injury by RWW larvae to rice roots resulted in altered oviposition by subsequent adults. The hypothesis for the second objective was that below‐ground RWW larval feeding would decrease plant suitability of rice to above‐ground RWW adults, resulting in decreased oviposition. 5. Results provided inconsistent support for the first hypothesis, indicating that responses differed among combinations of variety and injury level. Conversely, consistent support for the second hypothesis was found, indicating that larval feeding on roots decreased suitability of rice plants for oviposition.     

Effects of plant diversity and structural complexity on parasitoid behaviour in a field experiment

1. In natural ecosystems, plants containing hosts for parasitoids are often embedded within heterogeneous plant communities. These plant communities surrounding host‐infested plants may influence the host‐finding ability of parasitoids. 2. A release‐recapture‐approach was used to examine whether the diversity and structural complexity of the community surrounding a host‐infested plant influences the aggregation behaviour of the leaf‐miner parasitoid Dacnusa sibirica Telenga and naturally occurring local leaf‐miner parasitoids. Released and locally present parasitoids were collected on potted Jacobaea vulgaris Gaertn.plants infested with the generalist leaf‐miner Chromatomyia syngenesiae Hardy. The plants were placed in experimentally established plant communities differing in plant diversity (1–9 species) and habitat complexity (bare ground, mown vegetation, and tall vegetation). Additionally, parasitoids were reared out from host mines on the trap plants. 3. Plant diversity did not influence the mean number of recaptured D. sibirica or captures of other locally present parasitoids but the number of recaptured parasitoids was influenced by habitat complexity. No D. sibirica parasitoids were recaptured in the bare ground plots or plots with mown vegetation. The mean number of recaptured D. sibirica generally increased with increasing complexity of the plant community, whereas locally present parasitoids were captured more frequently in communities with more bare ground. There was a unimodal relationship between the number of reared out parasitoids and diversity of the surrounding vegetation with the highest density of emerged parasitoids at intermediate diversity levels. 4. The present study adds to the thus far limited body of literature examining the aggregation behaviour of parasitoids in the field and suggests that the preference of parasitoids to aggregate in complex versus simple vegetation is association specific and thus depends on the parasitoid species as well as the identity of the plant community.     

Contemporary evolution of host plant range expansion in an introduced herbivorous beetle Ophraella communa

Host range expansion of herbivorous insects is a key event in ecological speciation and insect pest management. However, the mechanistic processes are relatively unknown because it is difficult to observe the ongoing host range expansion in natural population. In this study, we focused on the ongoing host range expansion in introduced populations of the ragweed leaf beetle, Ophraella communa, to estimate the evolutionary process of host plant range expansion of a herbivorous insect. In the native range of North America, O. communa does not utilize Ambrosia trifida, as a host plant, but this plant is extensively utilized in the beetle's introduced range. Larval performance and adult preference experiments demonstrated that native O. communa beetles show better survival on host plant individuals from introduced plant populations than those from native plant populations and they also oviposit on the introduced plant, but not on the native plant. Introduced O. communa beetles showed significantly higher performance on and preference for both introduced and native A. trifida plants, when compared with native O. communa. These results indicate the contemporary evolution of host plant range expansion of introduced O. communa and suggest that the evolutionary change of both the host plant and the herbivorous insect involved in the host range expansion.     

The piercing-sucking herbivores Lygus hesperus and Nezara viridula induce volatile emissions in plants

Plant volatiles induced by herbivory are often used as olfactory cues by foraging herbivores and their natural enemies, and thus have potential for control of agricultural pests. Compared to chewing insects and mites, little is known about plant volatile production following herbivory by insects with piercing-sucking mouthparts. Here, we studied factors (insect life stage, gender, the role of salivary glands, and type of bioassay used for volatile induction) that influence the induction of plant volatiles by two agriculturally important hemipterans, Lygus hesperus and Nezara viridula. Feeding on intact cotton by virgin females of L. hesperus induced 2.6-fold greater volatile response compared to that induced by mated females, possibly due to increased feeding activity by virgin females. This plant volatile response was associated with elicitors present in the insect's salivary glands as well as to the degree of mechanical injury. Feeding injury by N. viridula females also increased volatile emissions in intact maize by approximately 2-fold compared to control plants. Maize seedlings injured by N. viridula emitted higher amounts of the monoterpene linalool, the sesquiterpenes (E)-beta-caryophyllene, alpha-trans-bergamotene, and (E,E)-beta-farnesene, and the homoterpene (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene, but not amounts of green leaf volatiles, compared to uninjured plants. Emissions from intact maize injured by adult males were lower than those emitted by adult females of the same age and did not differ from those emitted by uninjured plants. Similarly, feeding by virgin female N. viridula followed by excision led to 64% higher quantities of volatiles compared to untreated plants. Volatile emission in excised plants, however, was considerably greater than in intact plants, suggesting that careful consideration must be given to bioassay design in studies of herbivore-induced plant volatiles. Salivary gland extracts of N. viridula led to sesquiterpene emissions approximately 2.5-fold higher than for controls, although no significant differences were observed for green leaf volatiles, monoterpenes, and homoterpenes. These results indicate that L. hesperus and female N. viridula feeding induce volatile production in plants, and that volatile production is affected by gender and life stage of the bug. Although oviposition and mechanical injury by stylets may increase release of volatiles, elicitors from salivary glands of L. hesperus and N. viridula also seem to play a role in the emission of plant volatiles.     

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.