Xylem Feeding Research Articles

Lethal and sublethal effect of heat shock on Phenacoccus solenopsis Tinsley (Hemiptera: Pseudococcidae).

Temperature is an important abiotic environmental factor, and is responsible for various kinds of behavioral and physiological changes in living organisms. Induced heat shock is associated with feeding behaviour, reproduction and reactive oxygen species (ROS) generation that causes oxidative damage. In this experiment, we examined the lethal and sublethal effects of heat shock on reproduction, feeding behaviour and antioxidant enzymes, including catalase (CAT), superoxide dismutase (SOD) and peroxidases (POD) in P. solenopsis. Results showed that males were highly susceptible to heat shock treatments than females, as LTemp50 values were 43.8°C for males and 45.11°C for females. Heat shock events non-significantly affected the fecundity in female only treated adults and significantly affected the both sexes heat treated adults, it increased the xylem feeding duration, percentage of xylem feeding adults and reduce the phloem feeding duration and percentage of phloem feeding adults. Similarly it alter the antioxidant enzymes activities, an increase of CAT, SOD and POD activities were noticed in response to highest intensity of heat shock while a reduction of CAT and SOD activity were noticed in response to lowest intensity of heat shock compared to control (30°C). These results suggest that heat shock may result in loss of body water and induce oxidative stress in P. solenopsis. However, antioxidant enzymes play a significant role in overcoming the oxidative damage.

Feeding Behavior of Asian Citrus Psyllid [Diaphorina citri (Hemiptera: Liviidae)] Nymphs and Adults on Common Weeds Occurring in Cultivated Citrus Described Using Electrical Penetration Graph Recordings.

Asian citrus psyllid, Diaphorina citri, transmits Candidatus Liberibacter asiaticus (CLas), the putative causal agent of Huanglongbing disease. Although they primarily feed on the phloem of Citrus and related plants, when grove or host conditions are unfavorable, D. citri may be able to use weed species as alternate food sources for survival. To explore this possibility, electrical penetration graph (EPG) recordings (18 h) were performed to investigate the feeding behavior of psyllid adults and nymphs on three common south Florida weeds (Bidens alba, Eupatorium capillifolium, and Ludwigia octovalvis). EPG recordings revealed that the proportion of time spent by D. citri feeding on xylem was similar on all tested weed species (19%–22%) and on the positive control (20%), the preferred host, Citrus macrophylla. Very little to no phloem feeding was observed on weed species by either nymphs or adults. Histological studies using epifluorescence microscopy showed that salivary sheaths were branched and extended into xylem of weed species, whereas they ended in phloem on citrus plants. No choice behavioral assays showed that adults can obtain some nutrition by feeding on weed species (xylem feeding) and they may be able to survive on them for short intervals, when host conditions are unfavorable.

Hexaacetyl-chitohexaose, a chitin-derived oligosaccharide, transiently activates citrus defenses and alters the feeding behavior of Asian citrus psyllid

Plants have a perception system triggered by pathogen and pest signals to initiate defense. These signals include evolutionarily conserved molecules from microbes and insects termed pathogen/herbivore-associated molecular patterns (PAMPs/HAMPs). Here we showed that hexaacetyl-chitohexaose (HC), an oligosaccharide from chitin, a structural component in insect exoskeletons and fungi cell walls, upregulated defense-associated genes WRKY22, GST1, RAR1, EDS1, PAL1 and NPR2, and downregulated ICS1 at 1 h after HC treatment in Sun Chu Sha mandarin leaves. The effect was transient as defense gene transcriptional changes were not observed at 18 h after the treatment. Electrical penetration graph (EPG) recordings were used to study the feeding behavior of Asian citrus psyllid (ACP) following the HC treatment. ACP is the hemipteran vector of Candidatus Liberibacter asiaticus (CLas), the pathogen associated with huanglongbing (HLB). Adult ACP displayed reduced intercellular probing, reduced xylem feeding count and duration, and increased non-probing activity on HC-treated citrus compared to controls. During an 18-h recording, percentage for total duration of xylem ingestion, phloem ingestion, intercellular probing were lower, and the percentage of non-probing behavior was higher in HC-treated leaves than in controls. In host-selection behavior studies, HC treatment did not alter the attractiveness of citrus leaves under light or dark conditions. In addition, ACP feeding on HC-treated leaves did not show differences in mortality for up to 10 day of exposure. In summary, we report that HC induced a transient defense in citrus and an inhibitory effect on ACP feeding but did not affect host selection or the insect fitness under the tested conditions.

Match and mismatch between dietary switches and microbial partners in plant sap-feeding insects.

Some animal groups associate with the same vertically transmitted microbial symbionts over extended periods of evolutionary time, punctuated by occasional symbiont switches to different microbial taxa. Here we test the oft-repeated suggestion that symbiont switches are linked with host diet changes, focusing on hemipteran insects of the suborder Auchenorrhyncha. These insects include the only animals that feed on plant xylem sap through the life cycle, as well as taxa that feed on phloem sap and plant parenchyma cells. Ancestral state reconstruction provides strong statistical support for a xylem feeding auchenorrhynchan ancestor bearing the dual symbiosis with the primary symbiont Sulcia (Bacteroidetes) and companion symbiont 'β-Sym' (β-proteobacteria). We identified seven dietary transitions from xylem feeding (six to phloem feeding, one to parenchyma feeding), but no reversions to xylem feeding; five evolutionary losses of Sulcia, including replacements by yeast symbionts, exclusively in phloem/parenchyma-feeding lineages; and 14-15 losses of β-Sym, including nine transitions to a different bacterial companion symbiont. Our analysis indicates that, although companion symbiont switching is not associated with shifts in host diet, Sulcia is probably required for xylem-feeding. Furthermore, the ancestral auchenorrhynchan bearing Sulcia and β-Sym probably represents the sole evolutionary origin of xylem feeding in the animal kingdom.

Lethal and Sublethal Effects of Neonicotinoid Insecticides on the Adults of Phenacoccus solenopsis (Hemiptera: Pseudococcidae) on Tomato Plants.

Acetamiprid and imidacloprid are two important neonicotinoid insecticides that are widely utilized under field conditions for the management of sucking insect pests, including the solenopsis mealybug Phenacoccus solenopsis Tinsley (Hemiptera: Pseudococcidae). Although some information is available regarding their lethal effects, nothing is currently known about the sublethal effects of these insecticides. We, therefore, performed a series of experiments to test the lethal and sublethal effects of these chemicals on oviposition duration and fecundity. We also assessed sublethal effects on feeding behavior using the electrical penetration graph (EPG) technique. The results of this study reveal that acetamiprid toxicity is higher than imidacloprid and that both insecticides have negative effects on the oviposition, fecundity, and feeding behavior of P. solenopsis when applied at sublethal dosages. These chemicals also significantly reduce oviposition duration and fecundity and significantly prolong nonprobing duration, increase penetration problems, and reduce phloem and xylem feeding activities when compared with adults exposed to just water. No significant differences were detected in all waveform durations and events when adults previously exposed to foliage treated with each of these two insecticides were compared. The results of this study, therefore, suggest that both insecticides are capable of protecting crops from mealybug damage by not only killing these pests directly but also reducing their fecundity and inhibiting feeding behaviors when applied at sublethal dosages.

Rice dwarf virus infection alters green rice leafhopper host preference and feeding behavior.

Host plants, pathogens and their herbivore vectors systems have complex relationships via direct and indirect interactions. Although there are substantial gaps in understanding these systems, the dynamics of the relationships may influence the processes of virus transmission and plant disease epidemics. Rice dwarf virus (RDV) is mainly vectored by green rice leafhoppers (GRLHs), Nephotettix cincticeps (Uhler) (Hemiptera: Cicadellidae) in a persistently circulative manner. In this study, host plant selection preferences of non-viruliferous and viruliferous (carrying RDV) GRLHs between RDV-free and RDV-infected plants were tested. Non-viruliferous GRLHs preferred RDV-infected rice plants over RDV-free rice plants, and viruliferous GRLHs preferred RDV-free rice plants over RDV-infected rice plants. In odor selection preference bioassay using a four-field olfactometer, non-viruliferous GRLHs preferred odors of RDV-infected rice plants over healthy rice and viruliferous GRLHs preferred odors of RDV-free rice plants over RDV-infected ones. In 6 h plant penetration behavior bioassay using electrical penetration graphs, non-viruliferous GRLHs spent shorter time in non-penetration and much longer time in xylem feeding on RDV-infected, compared to RDV-free rice plants. Viruliferous GRLHs exhibited more salivation and stylet movement on RDV-free rice plants than on RDV-infected rice plants. We infer from these findings that RDV influences these vector behaviors by altering host plant physiology to promote viral transmission.

Feeding behaviors of Cacopsylla pyricola (Hemiptera: Psyllidae) using electrical penetration graphs (EPGs)

The pear psylla, Cacopsylla pyricola, is a very small sap-feeding insect of many commercial pear varieties that could be considered the most serious insect pest of pear. Detailed information on plant penetration activities of the pear psylla is essential to study its feeding behavior used to evaluate resistant traits to chemical control. The application of the electrical penetration graph technique (EPG) could provide a relevant insight into the nature of this resistance. EPG waveforms of C. pyricola were characterized on the basis of amplitude, frequency, voltage level, and electrical origin. Feeding behaviors of C. pyricola were recorded and analyzed by EPG analysis. During EPG monitoring, waveform PA occurred at the start of stylet penetration of pear leaf epidermal cell. Waveform PB followed, in which stylet secreted saliva was observed. Waveforms PC1 and PC2 involved penetrating and sucking behaviors in parenchyma cells and vascular parenchyma, respectively. In addition, waveform PC1 represented salivation into bundle sheath cells and ingestion from parenchyma. Otherwise, behaviors of salivation into phloem and ingestion from phloem produced waveforms PE1 and PE2, respectively. On the other hand, ingestion from xylem tissues showed waveform PG. Among the feeding patterns of C. pyricola described above, phloem feeding patterns occurred most frequently, followed by xylem feeding and parenchyma penetration patterns in descending order.

Plant penetration activities by the flatid planthopper Metcalfa pruinosa (Hemiptera: Fulgoroidea): an electrical penetration graph‐histology analysis

AbstractThe citrus flatid planthopper, Metcalfa pruinosa Say (Hemiptera: Flatidae), is a very polyphagous native insect in North America and currently a serious pest in Europe and South Korea. To understand the feeding behaviour of M. pruinosa, stylet penetration behaviour of M. pruinosa was investigated with an electrical penetration graph (EPG) system. This study reports seven EPG waveforms related to M. pruinosa feeding behaviour: np (no stylet penetration), Mp1 (initiation of stylet penetration), Mp2 (stylet movement and salivation), Mp4 (phloem feeding), Mp4‐H (honeydew excretion), Mp5 (xylem feeding) and Mp6 (unknown). To determine respective feeding behaviour related to the Mp4 and Mp5 waveforms, stylets were cut with a laser beam, and the location of the stylet tip within plant tissue was examined. We found plant sap was exuded from the severed stylets only when the Mp4 waveform was observed, suggesting phloem sap ingestion. The stylet tip was located in the xylem region, indicating xylem‐feeding activity, when the Mp5 waveform was observed. The analysis of 24 different EPG parameters suggests that M. pruinosa stylets reached the vascular bundle of a plant within ca. 5 min and spend ca. 70% of the time feeding on xylem and phloem feeding. This is the first study that reports seven distinctive EPG waveforms with respect to the feeding behaviour of M. pruinosa which could help determine host specificity and host plant susceptibility.

Varied responses by yeast-like symbionts during virulence adaptation in a monophagous phloem-feeding insect

This study examines the three-way interaction between symbionts, insect herbivores and their host plants during adaptation to resistant crop varieties. We conducted a long-term selection study (20 generations of continuous rearing) with a monophagous phloem-feeder, the brown planthopper [Nilaparvata lugens (Stal)], on several resistant rice (Oryza sativa L.) varieties. Planthopper fitness and the abundance of yeast-like symbionts (YLS) were monitored throughout the selection process. N. lugens populations collected from six regions in the Philippines adapted to the resistant varieties as noted by increasing body size and increased egg-laying. Adaptation was partially through physiological and behavioral changes apparent during feeding: Planthoppers on resistant plants had relatively high levels of xylem feeding compared with planthoppers on susceptible plants. YLS densities were highly dependent on the host rice variety. However, there were no consistent trends in YLS density during host plant switching and virulence adaptation: Compared to densities in planthoppers on the standard susceptible variety Taichung Native 1 (TN1), YLS densities were consistently higher on PTB33 (resistant), similar on IR62 (resistant) and IR65482 (moderately resistant) but lower on IR22 (susceptible). Furthermore, YLS densities often remained the same despite improved planthopper fitness over generations. Our results do not support the hypothesis that changes in YLS density mediate planthopper adaptation to resistant varieties. However, slight reductions in YLS densities toward the end of selection on TN1, IR22 and IR62 may indicate that YLS have lower functional significance where varieties and environmental conditions are constant between generations.

A terrestrial invader threatens a benthic community: potential effects of hemlock woolly adelgid-induced loss of eastern hemlock on invertebrate shredders in headwater streams

The invasion of non-native species is considered among the most important causes of the loss of native biota. An example of a devastating exotic pest impacting eastern forests is the hemlock woolly adelgid (HWA) (Adelges tsugae), a xylem feeding insect native to Asia that feeds on all ages and species of hemlock (Tsuga spp.), and has the capacity to functionally eliminate eastern hemlock (T. canadensis) from the landscape. Eastern hemlock is an important component of forested riparian communities in southern and central Appalachia where it maintains stable stream conditions important for aquatic biota. We investigated differences between eastern hemlock and deciduous dominated headwater streams in eastern Kentucky to determine the extent to which HWA invasion may affect benthic invertebrate shredders. Shredders form the basis of a resource chain within headwater streams, thus they are invaluable links in stream-riparian interactions. We found that eastern hemlock contributed energy in the form of leaf litter to the riparian zone at a consistent rate throughout the growing season in contrast to the autumnal pulse of deciduous material. We also detected significant vegetation by season interactions for shredders which suggest that some shredders are more abundant in eastern hemlock streams than in their deciduous counterparts during the summer. This could be linked to the consistent addition of food resources that eastern hemlock provides. Our results indicate that stream communities will likely change following adelgid associated mortality of eastern hemlock in headwater stream riparian zones.

Lietuvoje aptikti vabzdžiai, galimi fitoplazmų ‘Candidatus Phytoplasma asteris’ ir ‘Ca. Phytoplasma pruni’ giminingų kamienų pernešėjai

Phytoplasma strains affiliated with groups 16SrI, 16SrIII, 16SrV and 16SrXII have been found in Lithuania, but their insect vectors in the country have not been determined. ‘Candidatus Phytoplasma asteris’ and ‘Ca. Phytoplasma pruni’-related phytoplasma strains were identified in five leafhopper species and three spittlebug species occurring in Lithuania. The occurrence in Lithuania of Anaceratagallia ribauti, reported as a vector of stolbur phytoplasma (‘Ca. Phytoplasma solani’, subgroup 16SrXII-A) elsewhere in Europe, provokes the question of whether this phytoplasma may be present but yet undetected in Lithuania. The finding of subgroup 16SrI-C strains in A. ribauti, Aphrodes sp., Macrosteles sexnotatus and Euscelis incisus; 16SrI-B strains – in Aphrophora alni, Aphrodes sp. and Cicadella viridis; 16SrIII-B strains – in A. ribauti; and 16SrIII-P – in Aphrodes sp. and E. incisus is consistent with the hypothesis that these leafhopper species are vectors of the respective phytoplasmas in Lithuania. Transmission trials were initiated based on these results. Results thus far have revealed that Lithuanian biotype of E. incisus is capable of vectoring subgroup 16SrI-C phytoplasma strains that are found in Lithuania. The presence of diverse phytoplasmas in three spittlebug species, Aphrophora alni, Lepyronia coleopterata and Philaenus spumarius, indicates that these xylem feeders actually ingested phloem sieve cell contents during feeding on phytoplasmainfected plants, bringing into focus the question of whether some of such xylem feeders might act as occasional vectors of phytoplasmas.

EPG를 이용한 담배가루이(Bemisia tabaci)의 섭식행동과 Cyantraniliprole에 대한 섭식변화

Feeding behaviors of the sweet potato whitefly, Bemisia tabaci, and changing of their feeding behaviors were recorded and analyzed with an Electrical Penetration Graph (EPG) analysis against cyantraniliprole. The characteristic patterns of feeding behaviors investigated were as follows; NP (non probing), pathway phase waveform, waveform of feeding xylem. In results, B. tabaci did not almost feeding the phloem. And It was not appeared potential drop during recording time. It was predominantly indicated pathway activity waveform and waveform of xylem feeding. After treatment of cyantraniliprole with recommended concentrations to tomato plants, EPG waveforms were recorded during 3 hours. Cyantraniliprole treatment showed longer time to first feeding xylem than untreated (P=0.043). It was showed shorter duration of first feeding xylem than untreated (P=0.017). And it was showed longer NP (non-probing) time than untreated. Consequently, It was supposed that cyantraniliprole was effect of anti-feeding or avoidance to B. tabaci.

Mathematical modeling of Glassy-winged sharpshooter population

Pierce's disease (PD) is a fatal disease of grapevines which results from an infection by the plant pathogen Xyllela fastidiosa. This bacterium grows in the xylem (water-conducting) vessels of the plant blocking movement of water. PD can kill vines in one year and poses a serious threat to both the California and the expanding Texas wine industries. Bacteria are vectored from one vine to the next by a number of xylem feeding insect species. Of these, the Glassy-winged Sharpshooter (GWSS) is considered to be the primary xylem feeding insect in Texas vineyards. An extensive database of the xylem-feeding population frequencies was collected by USDA-APHIS for Texas vineyards over multiple years. This project focused on a subset of data, GWSS frequencies within 25 vineyards in Edwards Plateau located in central Texas. The proposed model investigates the natural population dynamics and the decline in GWSS, likely the result of pest management campaigns on the insects within the region. The model is a delay Gompertz differential equation with harvesting and immigration terms, and we use the data to estimate the model parameters.

EPG technique as a tool to reveal host plant acceptance by xylem sap‐feeding insects

AbstractThe discovery of a new exotic insect herbivore triggers responses from biosecurity agencies, one of which is the decision of whether or not to attempt eradication. Rapid determination of the host range of the new invader is necessary, but when sap‐sucking insects are first collected from plants, the lack of visible signs of feeding damage makes it difficult to determine their host status. We investigated the Electrical Penetration Graph (EPG‐DC) technique as tool to assess host range of a xylem sap‐feeding invader, using Carystoterpa fingens (Hemiptera: Aphrophoridae), a New Zealand endemic xylem feeder, as a model insect. Real‐time probing and feeding events over a 12‐h recording period were compared for adult C. fingens on 18 plant species. Hebe azure, a known host, was designated the ‘reference plant species’ against which events on all other plants were statistically compared. EPG waveforms were categorized on their amplitude, frequency, voltage and electrical origin, and six parameters (time taken to first probe, time to first xylem ingestion from first probe, total probing time, number of xylem‐ingesting events, duration of the longest xylem‐ingesting event and total xylem ingestion time) were measured. The total xylem ingestion period (i.e. the actual feeding period) on each plant species expressed as a percentage of total probing time was considered the best parameter for comparing the host status of plants with H. azure. Although the EPG data overestimated the actual host range of C. fingens, we consider that they provided a reasonable first guide to the potential host status of the unknown plants, and so might usefully be used to rapidly assess whether a plant from which a new invader was collected was a host, or whether the association was merely incidental.

Seasonal Abundance of Draeculacephala minerva and Other Xylella fastidiosa Vectors in California Almond Orchards and Vineyards

Almond leaf scorch (ALS) disease is caused by the bacterium Xylella fastidiosa and transmitted by xylem-feeding insects. Reports of increased incidence of ALS-diseased trees in California prompted surveys in three almond [Prunus dulcis (Mill.) D. A. Webb]-growing regions, from June 2003 to September 2005, to determine insect vector species composition and abundance. For comparison, sampling in and near vineyards in the San Joaquin Valley, California, also was completed. Sampling in or near almond orchards collected >42,000 Cicadomorpha of which 4.8% were xylem feeders, including 1912 grass sharpshooter, Draeculacephala minerva Ball; five Xyphon fulgida Nottingham; and a single spittlebug, Philaenus spumarius L. The most abundant vector was D. minerva. Season-long sampling indicated that D. minerva was a year-round resident in and/or near almonds in the Sacramento Valley, but not in the San Joaquin Valley. Similarly, D. minerca was rare in vineyards in the San Joaquin Valley, but was abundant in irrigated pastures near vineyards. D. minerva was most frequently collected along orchard margins, and peak densities were observed in summer, the period of time when bacterial titers are reported to increase in infected trees. Screening of D. minerva for presence of X.fastidiosa found that 1.1% of insects collected near almond orchards and 4.5% of insects collected from pastures tested positive. The X. fastidiosa subspecies and genotype detected in insects collected from orchards matched those collected from ALS-diseased almond trees in the same orchard. Of the few X. fulgida and P. spumarius collected, none tested positive for X. fastidiosa. Results are discussed with respect to X. fastidiosa vector control and detection methods.

Insect herbivore mortality is increased by competition with a hemiparasitic plant

Summary 1. Individual plants are often simultaneously consumed by many different organisms and mediate important indirect interactions between their consumers, even when their consumers are phylogenetically distant and feed on different parts of the plant. 2. We examined the effects on two xylem feeders of sharing a plant host in greenhouse pot trials and outdoor mesocosms. The spittle bug Neophilaenus lineatus, which feeds on grass shoots, and a hemiparasitic angiosperm, Rhinanthus minor, which attaches to the roots, were added separately or together on grass host plants growing in pots or in model grassland swards. 3. We found strong asymmetric competition between the hemiparasite and the insect. In greenhouse experiments, sharing a grass host with R. minor significantly increased mortality of N. lineatus, but R. minor performance was unaffected. 4. Mortality of N. lineatus was also increased on swards containing R. minor in model communities grown in the field. R. minor was more sensitive to sharing a host with N. lineatus in these communities, and experienced a modest (non‐significant) reduction in flower production. 5. The hemiparasite was the stronger competitor for xylem resources, possibly because of its ability to extract resources from the plant roots, and because of its significant ability to appropriate the xylem stream via its strong transpirational pull. 6. We believe these are the first experiments to demonstrate competition between insect herbivores and hemiparasitic plants mediated via a shared host plant. The consequences for the invertebrate population were potentially highly significant and we propose that future research should further explore the role of this asymmetric relationship in structuring invertebrate communities.

Independence of Resistance in Brachiaria spp. to Nymphs or to Adult Spittlebugs (Hemiptera: Cercopidae): Implications for Breeding for Resistance

Both nymphal and adult spittlebugs (Hemiptera: Cercopidae) cause serious economic damage to susceptible brachiariagrass [genus Brachiaria (Trin.) Griseb], pastures in tropical America. Both life stages are xylem feeders: nymphs feed primarily on roots and stems, whereas the adults feed mainly on foliage. Numerous interspecific brachiariagrass hybrids with high levels of antibiosis resistance to nymphs of several important spittlebug species have been obtained. Recent studies revealed major inconsistencies between reaction to nymphs and reaction to adults on the same host genotype. Because both insect life stages can cause severe economic damage on susceptible brachiariagrass pastures, a cultivar development strategy must take into account resistance to both life stages. To assess the degree of association between resistance to spittlebug nymphs and to adult feeding, we tested 164 hybrids and six check genotypes for resistance to both life stages of three spittlebug species: Aeneolamia varia (F.), Aeneolamia reducta (Lallemand), and Zulia carbonaria (Lallemand). Most hybrids tested were classified as resistant to nymphs. On the contrary, for all three species, the overall mean damage score of the 164 hybrids did not differ from the mean score of the susceptible checks. None of the hybrids was classified as resistant to adult feeding damage. Correlations between percentage nymph survival and adult damage scores were consistently low (r = 0.0104-0.0191). Correlations between nymphal and adult damage scores were also low (0.109-0.271), suggesting that resistances to the different life stages are largely independent. Chi-square analyses comparing frequency distributions of responses of the 164 breeding hybrids to nymphs or adults confirmed essential genetic independence of these two traits. We conclude that attention to improving genetic resistance specifically to adult feeding damage is warranted.

Effects of sap-feeding insect herbivores on growth and reproduction of woody plants: a meta-analysis of experimental studies

The majority of generalisations concerning plant responses to herbivory are based on studies of natural or simulated defoliation. However, effects caused by insects feeding on plant sap are likely to differ from the effects of folivory. We assessed the general patterns and sources of variation in the effects of sap feeding on growth, photosynthesis, and reproduction of woody plants through a meta-analysis of 272 effect sizes calculated from 52 papers. Sap-feeders significantly reduced growth (-29%), reproduction (-17%), and photosynthesis (-27%); seedlings suffered more than saplings and mature trees. Deciduous and evergreen woody plants did not differ in their abilities to tolerate damage imposed by sap-feeders. Different plant parts, in particular below- and above-ground organs, responded similarly to damage, indicating that sap-feeders did not change the resource allocation in plants. The strongest effects were caused by mesophyll and phloem feeders, and the weakest by xylem feeders. Generalist sap-feeders reduced plant performance to a greater extent than did specialists. Methodology substantially influenced the outcomes of the primary studies; experiments conducted in greenhouses yielded stronger negative effects than field experiments; shorter (<12 months) experiments showed bigger growth reduction in response to sap feeding than longer experiments; natural levels of herbivory caused weaker effects than infestation of experimental plants by sap-feeders. Studies conducted at higher temperatures yielded stronger detrimental effects of sap-feeders on their hosts. We conclude that sap-feeders impose a more severe overall negative impact on plant performance than do defoliators, mostly due to the lower abilities of woody plants to compensate for sap-feeders' damage in terms of both growth and photosynthesis.

The considerable adult size variability in wood feeders is optimal

1. The life history of wood feeders was modelled in order to explain the multiseasonality of development and the great variability of adult size in this group.2. The model was parameterised with experimental bioenergetic and reproductive data for the xylem feeder Aredolpona rubra (Coleoptera: Cerambycidae).3. The length of the developmental period, which together with food quality directly determines adult size and indirectly determines the number of eggs laid, was optimised.4. The results show that multi‐seasonal larval development maximises fitness under conditions of low food quality, relatively low predation pressure, and the presence of hostile periods during the year.5. The variability of the number of seasons needed to complete development within a wood‐feeder population is a consequence of development time optimisation and the unavoidable extension of the egg‐laying period. These insects have an evolutionary dilemma: to eclose late in a given season at smaller size, bringing about low egg production and low offspring value, or to grow bigger to the next season, jeopardising their lives.6. The results of the model predict wood‐feeder developmental patterns that depend on the tree tissue inhabited.

Silicon‐augmented resistance of plants to herbivorous insects: a review

AbstractSilicon (Si) is one of the most abundant elements in the earth's crust, although its essentiality in plant growth is not clearly established. However, the importance of Si as an element that is particularly beneficial for plants under a range of abiotic and biotic stresses is now beyond doubt. This paper reviews progress in exploring the benefits at two‐ and three‐trophic levels and the underlying mechanism of Si in enhancing the resistance of host plants to herbivorous insects. Numerous studies have shown an enhanced resistance of plants to insect herbivores including folivores, borers, and phloem and xylem feeders. Silicon may act directly on insect herbivores leading to a reduction in insect performance and plant damage. Various indirect effects may also be caused, for example, by delaying herbivore establishment and thus an increased chance of exposure to natural enemies, adverse weather events or control measures that target exposed insects. A further indirect effect of Si may be to increase tolerance of plants to abiotic stresses, notably water stress, which can in turn lead to a reduction in insect numbers and plant damage. There are two mechanisms by which Si is likely to increase resistance to herbivore feeding. Increased physical resistance (constitutive), based on solid amorphous silica, has long been considered the major mechanism of Si‐mediated defences of plants, although there is recent evidence for induced physical defence. Physical resistance involves reduced digestibility and/or increased hardness and abrasiveness of plant tissues because of silica deposition, mainly as opaline phytoliths, in various tissues, including epidermal silica cells. Further, there is now evidence that soluble Si is involved in induced chemical defences to insect herbivore attack through the enhanced production of defensive enzymes or possibly the enhanced release of plant volatiles. However, only two studies have tested for the effect of Si on an insect herbivore and third trophic level effects on the herbivore's predators and parasitoids. One study showed no effect of Si on natural enemies, but the methods used were not favourable for the detection of semiochemical‐mediated effects. Work recently commenced in Australia is methodologically and conceptually more advanced and an effect of Si on the plants' ability to generate an induced response by acting at the third trophic level was observed. This paper provides the first overview of Si in insect herbivore resistance studies, and highlights novel, recent hypotheses and findings in this area of research. Finally, we make suggestions for future research efforts in the use of Si to enhance plant resistance to insect herbivores.