Growth and defence in young pine and spruce and the expression of resistance to a stem-feeding weevil

Defence in young trees has been much less studied than defence in older ones. In conifers, resin within ducts in bark is an important quantitative defence, but its expression in young trees may be influenced by developmental or physical constraints on the absolute size of the resin ducts as well as by differential allocation of resources to growth and resin synthesis. To examine these relationships, we used nitrogen fertilisation of 1- and 2-year-old pine and spruce to produce trees of different sizes and measured the effect on the number and size of resin ducts and the amount of resin they contained. All of these variables tended to increase with stem diameter, indicating a positive relationship between resin-based defence and growth of 1- and 2-year-old trees. In pine, however, the mass of resin flowing from severed ducts was much lower relative to duct area in 1- than in 2-year-old trees, suggesting that the older trees allocated a higher proportion of the carbon budget to resin synthesis. Resin-based defence in 1-year-old pines appears to be both positively related to growth and resource limited. In spruce, resin production was generally lower, and age-related differences were not observed, suggesting that resin-based defence is less important in this species. Bio-assays of 2-year-old trees with the pine weevil, Hylobius abietis, emphasised the importance of resin as a defence against this bark feeding insect. Nitrogen fertilisation had a limited influence on resistance expression. One-year-old trees remained susceptible because of their small size, low resin production and limited response to fertilisation. The strong growth response of 2-year-old trees to fertilisation increased resin-based defence, but most spruce trees remained susceptible, while most pines were resistant at all levels of fertilisation.     

Reduced oviposition by Diaprepes abbreviatus (Coleoptera: Curculionidae) and growth enhancement of citrus by Surround particle film

Regularly applied sprays of a particle film, Surround WP, greatly enhanced the growth of citrus trees on a poorly drained Winder soil at Fort Pierce, FL. After 3 yr of applications every 3 or 4 wk, Surround-treated trees had at least 5 times the mass, 6 times the canopy volume, and approximately 4 times the cross-sectional area of the tree stems at the graft union compared with untreated trees. The larger Surround-treated trees attracted a higher number of adult weevil Diaprepes abbreviatus (L.) and to a lesser extent citrus root weevil, Pachnaeus litus (Germar), per tree, but there was an equivalent number of egg masses per tree compared with the control trees. The number of egg masses per female weevil oviposited on Surround-treated trees was significantly less than either the control trees or trees treated biannually with an entomopathogenic nematode, BioVector. The number of larvae per tree recovered from the roots of excavated trees was greater from trees treated with Surround once every 3 wk compared with control trees. The data suggest that Surround particle film greatly enhanced the growth of citrus trees grown in a poorly drained soil. The reduction in oviposition by D. abbreviatus was insufficient to significantly reduce the number of root weevil larvae per tree feeding on the roots. However, the more vigorous trees resulting from Surround applications may be more resistant or tolerant to root weevil feeding.     

Feeding behaviour and efficiency of banana weevil (Cosmopolites sordidus) larvae on banana cultivars of varying resistance levels

Nutrient composition, secondary plant substances and other host quality characteristics govern resistance and influence herbivore feeding. The current study was carried out to compare aspects of feeding behaviour and food utilization of banana weevil (Germar) (Coleptera: Curculionidae) larvae feeding on banana cultivars with different levels of resistance. In the first study, penetration times of larvae into tissue of seven banana cultivars (two susceptible, three intermediate and four resistant) were compared. Day‐old larvae were placed on corm or pseudostem tissue of these cultivars and time taken to penetrate the tissue was recorded. Larvae generally took longer to tunnel into resistant cultivars, although Yangambi Km5 (resistant) did not fit this pattern. In the second set of studies, food consumption and growth of larvae were compared on cultivars: (i) Kisubi (resistant) vs. Atwalira (susceptible); and (ii) Mbwazirume (intermediate) vs. Kayinja (resistant). Day‐old larvae were offered weighed pieces of corm that were changed every 4–7 days. At each corm change, weights of the larvae, corms and frass were measured. Changes in corm and larval weight were calculated to determine larval growth and amount of tissue consumed. Larvae feeding on resistant cultivars had lower consumption and growth rates and longer developmental duration. It was concluded that low food consumption rates contribute to delayed development of larvae feeding on resistant cultivars. Differences between resistant and susceptible cultivars in food utilization may be of limited consequence for the cultivars tested. This is one step in identifying the factors influencing weevil development rates in cultivars of different resistance levels. If these factors are identified, they can be targeted in breeding for resistance.     

Bioefficacy and mode-of-action of some limonoids of salannin group fromAzadirachta indica A. Juss and their role in a multicomponent system against lepidopteran larvae

Biological activities of the salannin type of limonoids isolated fromAzadirachta indica A. Juss were assessed using the gram pod borerHelicoverpa armigera (Hubner) and the tobacco armywormSpodoptera litura (Fabricius) (Lepidoptera: Noctuidae). Inhibition of larval growth was concomitant with reduced feeding by neonate and third instar larvae. All three compounds exhibited strong antifeedant activity in a choice leaf disc bioassay with 2.0, 2.3 and 2.8 (μ/cm² of 3-O-acetyl salannol, salannol and salannin, respectively deterring feeding by 50% inS. litura larvae. In nutritional assays, all three comounds reduced growth and consumption when fed to larvae without any effect on efficiency of conversion of ingested food (ECI), suggesting antifeedant activity alone. No toxicity was observed nor was there any significant affect on nutritional indices following topical application, further suggesting specific action as feeding deterrents. When relative growth rates were plotted against relative consumption rates, growth efficiency of theH. armigera fed diet containing 3-O-acetyl salannol, salannol or salannin did not differ from that of starved control larvae (used as calibration curve), further confirming the specific antifeedant action of salannin type of limonoids. Where the three compounds were co-administered, no enhancement in activity was observed. Non-azadirachtin limonoids having structural similarities and explicitly similar modes of action, like feeding deterrence in the present case, have no potentiating effect in any combination.     

Population dynamics of the white pine weevil, Pissodes strobi, infesting jack pine, Pinus banksiana, in Ontario, Canada

1. Eight generations of white pine weevil, Pissodes strobi (Peck) , infesting the terminal shoots of young jack pine trees, were sampled as larvae, pupae, and adults between 1988 and 1995.
2. The density of adult weevils increased rapidly for the first 3 years of the study, then declined in the next 4 years. Between-generation rates of change in density of weevils were related to mean number of weevils produced per terminal shoot of the host tree. Net increases in population density followed years in which a relatively high number of weevils emerged per terminal shoot and vice versa.
3. The mean number of weevils emerging per terminal shoot was determined by survival of weevils between the end of the larval stage and the successful emergence of adult weevils.
4. There was no clear relationship between survival of the post-feeding stages of the weevil and rates of parasitism or bird predation on these same stages. There was, however, a strong negative relationship between survival of weevils within the terminal shoot and the abundance of the facultative dipteran predator Lonchaea corticis Taylor.
5. The relationship among generational rates of change in populations, survival, and predation was evident irrespective of the age of the trees infested, suggesting that population dynamics of the weevil may be influenced as much by local predation pressure as by physical and biotic changes associated with growth of the young host trees.     

Early-season aerial adult colonization and ground activity of pea leaf weevil (Coleoptera: Curculionidae) in pea as influenced by tillage system

The pea leaf weevil, Sitona lineatus (L.) (Coleoptera: Curculionidae), is an important pest of pea, Pisum sativum L., in northern Idaho. Previous research revealed greater relative pea leaf weevil abundance and feeding damage in peas grown using conventional-tillage compared with no-tillage practices. However, the effects of tillage practices on early season colonization and activity by the pea leaf weevil on pea are not fully understood. Aerial traps and pitfall traps were used to assess adult colonization and relative density of adult pea leaf weevil into conventional-tillage and no-tillage pea in northern Idaho during 2005 and 2006. Feeding damage to the crop also was evaluated. During both years, aerial traps captured more pea leaf weevil in May, when crop establishment and early growth occurred, than in later months. Significantly more adult pea leaf weevils were captured in aerial traps in conventional-tillage than in no-tillage plots in May of both years. Significantly more pea leaf weevil were captured in pitfall traps in conventional-tillage plots than in no-tillage plots during the period immediately after peak aerial adult colonization in late May and early June. Crop feeding damage was significantly greater in conventional-tillage than in no-tillage plots in late May and early June. The patterns suggest that more adult pea leaf weevil colonize conventional-tillage pea than no-tillage pea. Pea plants in conventional-tillage emerged earlier and were larger than those in no-tillage during the pea leaf weevil colonization period, possibly accounting for the differences in colonization rates. This leads to greater early season pea leaf weevil infestation of conventional-tillage plots at a critical period for pea development that might ultimately influence crop yield.     

Differential defoliation of Eucalyptus grandis arises from indiscriminant oviposition and differential larval survival

1 The influence of six open-pollinated families (OPFs) of Eucalyptus grandis on both the growth and development of larvae and the oviposition preference of a paropsine chrysomelid ( Paropsis atomaria ) was investigated. The OPFs had previously been identified as differing in their susceptibility to defoliation by P. atomaria in forestry progeny trials.
2 Oviposition preference for resistant and susceptible foliage was tested using binary choice tests. These tests did not demonstrate any significant preference for either resistant or susceptible open-pollinated material indicating that adult host preference for susceptible trees was not a likely cause of differential defoliation.
3 Quantification and analysis of growth and development parameters for all larval stages of P. atomaria showed that feeding on genetic material identified as resistant resulted in a significant reduction of relative growth rate of first instar larvae and an alteration to normal feeding behaviour. There was also a trend towards increased larval mortality on resistant E. grandis .
4 We argue that although the magnitude of these effects was minor, interactions with additional biotic and abiotic sources of mortality in the field have the potential, when magnified over successive generations, to result in significant variation in defoliation of host genotypes in the field.     

Moisture content and nutrition as selection forces for emerald ash borer larval feeding behaviour

1. The exotic phloem‐feeding emerald ash borer (EAB), Agrilus planipennis, has killed tens of millions of North American ash trees (Fraxinus) since its first detection in the U.S.A. in 2002. Ash trees are killed by larval feeding in the cambial region, which disrupts translocation of photosynthates and nutrients. 2. We observed that EAB larvae feed predominantly downwards in naturally grown green ash trees, a behaviour confirmed in greenhouse‐grown black ash seedlings. Furthermore, biomass of larvae feeding downwards was greater than that for larvae feeding upwards. 3. We sought to determine the relative importance of four selection forces (i.e. gravity, moisture content, plant defence, and nutrition) in driving this downward feeding behaviour in this study. The gravity and plant defence (i.e. polyphenols) hypotheses were ruled out because even when seedlings were grown upside down, more EAB larvae moved upwards (towards the root area), and phloem tissue below the feeding site contained higher concentrations of defensive compounds than that above the feeding site. 4. The moisture content hypothesis was supported as phloem moisture above the feeding site decreased to levels reducing survivorship and biomass but was unaffected below. The nutrition hypothesis was also supported as the levels of 11 amino acids (mostly essential amino acids) below the feeding site were greater than those above. Furthermore, growth of larvae reared on an artificial diet deficient in protein and amino acids was worse than larvae reared in diet with complete ingredient or diet deficient in either protein or amino acids. 5. We conclude that moisture content and nutrients are two selective forces for the downward feeding behaviour of EAB larvae.     

Physical defences wear you down: progressive and irreversible impacts of silica on insect herbivores

1. Silica in the leaves of grasses can act as a defence against both vertebrate and invertebrate herbivores. The mechanisms by which silica affects herbivore performance are not well characterized. Here we expose an insect herbivore Spodoptera exempta to high-silica diets and test two mechanisms by which silica has been proposed to act as a defence. First, that silica reduces the digestibility of leaves and second, that silica causes wear to insect mandibles, both of which could potentially impact on herbivore performance. 2. Silica reduced the efficiency with which S. exempta converted ingested food to body mass and the amount of nitrogen absorbed from their food, leading to reduced insect growth rates. The measure of how efficiently herbivores utilize digested food (ECD) was unaffected by silica. 3. These effects occurred even with short-term exposure to silica-rich diets, but they also increased markedly with the duration of exposure and affected late instars more than early instar larvae. This appears to be due to the progressive impacts of silica with longer exposure times and suggests that herbivores cannot adapt to silica defences, nor do they develop a tolerance for silica with age. 4. Exposure to silica-rich diets caused increased mandible wear in S. exempta. This effect was extremely rapid, occurring within a single instar, further reducing feeding efficiency and growth rates. These effects on insect growth and feeding efficiency are nonreversible, persisting after the herbivore has switched diets. Up to a third of this residual impact can be explained by the degree of mandible wear caused by previous silica-rich diets. 5. The impacts of silica on S. exempta larvae were progressive with exposure time and could not be compensated for, even by switching to a different diet. Thus, herbivores cannot easily adapt to physical defences such as silica, suggesting this defence will have major implications for herbivore fitness.     

A molecular and genomic reference system for conifer defence against insects

Insect pests are part of natural forest ecosystems contributing to forest rejuvenation but can also cause ecological disturbance and economic losses that are expected to increase with climate change. The white pine or spruce weevil (Pissodes strobi) is a pest of conifer forests in North America. Weevil–host interactions with various spruce (Picea) species have been explored as a genomic and molecular reference system for conifer defence against insects. Interactions occur in two major phases of the insect life cycle. In the exophase, adult weevils are free‐moving and display behaviour of host selection for oviposition that is affected by host traits. In the endophase, insects live within the host where mobility and development from eggs to young adults are affected by a complex system of host defences. Genetic resistance exists in several spruce species and involves synergism of constitutive and induced chemical and physical defences that comprise the conifer defence syndrome. Here, we review conifer defences that disrupt the weevil life cycle and mechanisms by which trees resist weevil attack. We highlight molecular and genomic aspects and a possible role for the weevil microbiome. Knowledge of this conifer defence system is supporting forest health strategies and tree breeding for insect resistance.     

Elm leaves ‘warned’ by insect egg deposition reduce survival of hatching larvae by a shift in their quantitative leaf metabolite pattern

Plants may take insect eggs on their leaves as a warning of future herbivory and intensify their defence against feeding larvae. Responsible agents are, however, largely unknown, and little knowledge is available on this phenomenon in perennial plants. We investigated how egg deposition affects the anti‐herbivore defence of elm against the multivoltine elm leaf beetle. Prior egg deposition caused changes in the quality of feeding‐damaged leaves that resulted in increased larval mortality and reduced reproductive capacity of the herbivore by harming especially female larvae. Chemical analyses of primary and secondary leaf metabolites in feeding‐damaged, egg‐free (F) and feeding‐damaged, egg‐deposited (EF)‐leaves revealed only small differences in concentrations when comparing metabolites singly. However, a pattern‐focused analysis showed clearly separable patterns of (F) and (EF)‐leaves because of concentration differences in especially nitrogen and phenolics, of which robinin was consumed in greater amounts by larvae on (EF) than on (F)‐leaves. Our study shows that insect egg deposition mediates a shift in the quantitative nutritional pattern of feeding‐damaged leaves, and thus might limit the herbivore's population growth by reducing the number of especially female herbivores. This may be a strategy that pays off in a long run particularly in perennial plants against multivoltine herbivores.     

The effects of feeding by the rice water weevil, Lissorhoptrus oryzophilus Kuschel, on the growth and yield components of rice, Oryza sativa

Abstract 1 The effects of feeding by larvae of the rice water weevil, Lissorhoptrus oryzophilus Kuschel, on the growth and yield components of rice, Oryza sativa, were evaluated using four varieties of rice, ‘Cocodrie’, ‘Cypress’, ‘Bengal’ and ‘XP1003’, over 2 years. 2 Both adults and larvae of L. oryzophilus feed on rice plants, but it is feeding by the larval stage that causes economic losses. Feeding by L. oryzophilus larvae resulted in extensive damage to root systems. Pruning of root systems resulted in a decrease in tiller number and shoot biomass of rice plants in the vegetative stage of growth. 3 Yield losses were due to a combination of decreases in panicle densities, numbers of grains per panicle, and grain weights. Decreases in panicle densities were a direct result of the reductions in tiller numbers. Reductions in numbers of grains per panicle and grain weights probably resulted from decreases in shoot biomass. 4 Injury by rice water weevil larvae is chronic. The tillering stage of rice suffered the majority of weevil damage, but the growth effects were not manifested until later.     

Walnut development affects chemical composition and codling moth performance

1 We investigated for early and late blooming walnut cultivars in California whether variation in nut phenology resulted in differences in nutritional quality and whether this, in turn, affected the performance of the codling moth, Cydia pomonella (L.), and the extent of nut damage. 2 Mid‐season, during the period of nut growth, nuts from the early cultivars were larger than those from the late cultivars and had higher nitrogen content in both husk and kernel tissue, while kernel phenolic content was significantly lower. No major differences were observed later in the season after nuts from all cultivars had reached their final size. 3 Throughout the season establishment of neonate larvae was highest on nuts from the early cultivars but this was only significantly so at the beginning of the third codling moth generation. During the second codling moth generation (mid season) relative growth rates of third‐instar larvae were significantly higher on early than on late cultivars. Nut damage in the field was also significantly greater on early than on late cultivars during generation 2, while no significant differences were observed during generation 3. 4 The data suggest that the variation in codling moth damage among walnut cultivars is related to bloom phenology due to the influence of nut phenology on larval performance.     

Spruce budworm growth,development and food utilization on young and old balsam fir trees

Laboratory rearing of spruce budworm, Choristoneura fumiferana, in conjunction with field rearing, gravimetric analyses, a transfer experiment, and foliage chemical analyses at six dates during the period of budworm feeding activity indicated that the age of balsam fir, Abies balsamea, trees (70-year-old mature trees or 30-year-old juvenile trees) affected tree suitability for the spruce budworm via the chemical profile of the foliage. Insects reared on old trees had greater survival and pupal weight, shorter development times, and caused more defoliation than those reared on young trees. Young trees were more suitable for the development of young larvae (instars 2–5), while old trees were more suitable for the development of older, sixth-instar larvae. These results were confirmed by the laboratory transfer experiment. Young larvae fed foliage from young trees had higher relative growth rates (RGR), digestibility (AD), and efficiency of conversion of ingested foliage (ECI) than those fed foliage from old trees. These differences appeared to be related to the high N:tannins ratio, and the high contents of P present in young trees during the development of the young larvae. Old larvae fed foliage from old trees had higher relative growth rates, relative consumption rates (RCR), and digestibility of the foliage than those fed foliage from young trees. The high digestibility of the foliage of old trees was compensated for by a lower efficiency of conversion of digested food (ECD), which in turn resulted in no significant effect of tree age on the efficiency of conversion of ingested foliage by old larvae. The low relative consumption rate of old larvae fed foliage from young trees appeared to be related to the low N:tannins ratio, and the high contents of bornyl acetate, terpinolene, and °-3-carene present in young trees during the budworm sixth instar. Variations in these compounds in relation to tree age may serve as mechanisms of balsam fir resistance to spruce budworm by reducing the feeding rate of sixth instar larvae.     

Microchemical analysis of laser-microdissected stone cells of Norway spruce by cryogenic nuclear magnetic resonance spectroscopy

Stone cells (sclereids) in Norway spruce (Picea abies) bark have been reported to be highly lignified tissues that are important in physical defence against bark beetle invasion. Microchemical analyses of the low-molecular weight compounds in the stone cells of Norway spruce were carried out using laser microdissection in combination with cryogenic nuclear magnetic resonance and mass spectrometry (LMD/NMR/MS). Two phenolic compounds, the stilbene astringin and the dihydroflavonol dihydroxyquercetin 3′-O-β-d-glucopyranoside, were identified indicating that stone cells are more than just repositories for lignin. Both of these compounds were also found to be present in other phloem tissue at a higher level than in the stone cells based on quantification by cryogenic ¹H NMR. Our results suggest that stone cells may be involved in chemical as well as physical defense against bark beetles and their associated microorganisms. This paper reports on the identification of secondary plant metabolites from a single laser-microdissected population of plant cells offering a sensitive new way to determine the chemical profile of specific plant cell types with a high degree of precision.     

The role of lignin as a defence against the spruce bark beetle Dendroctonus micans: effect on larvae and adults

Summary The role of lignin as a physical defence against Dendroctonus micans was investigated in laboratory feeding experiments. The effect of lignin is dose-dependent, reducing larval survival, growth rate, and weight, as well as affecting gallery construction. Adults lay fewer eggs in lignified bark and also tend to construct abnormal galleries. The distribution of lignin in trees suggests a role in defence against bark beetles that feed in the thicker bark on the lower bole.     

Effect of kaolin on fitness and behavior of Choristoneura rosaceana (Lepidoptera: Tortricidae) larvae

The mechanisms by which kaolin, a clay particle film formulation, affects the fitness and behavior of larvae of obliquebanded leafroller, Choristoneura rosaceana (Harris), were investigated. Feeding experiments tested kaolin as a physical barrier versus a physiological toxin for larvae that consumed kaolin applied either to apple (Malus spp.) leaves or mixed in artificial diet. Behavioral experiments tested the effects of kaolin applied to apple leaves on neonate dispersal and leaf rolling by third and fourth instars. When larvae fed on apple leaves sprayed with kaolin, mortality and time to pupation of larvae increased significantly, whereas pupal mass significantly decreased. When larvae consumed kaolin mixed into an artificial diet, however, the effects on mortality, pupation time, and pupal mass were negligible. There may be minor physiological effects from consumption because male time to pupation was delayed for larvae fed diets containing the highest concentration of kaolin. In behavioral experiments, neonate larvae dispersed more quickly off plants covered with kaolin than control plants, and kaolin delayed the construction of leaf shelters by third and fourth instars. We showed that the effects of kaolin on C. rosaceana larvae are primarily physical, causing changes in dispersal and rolling behaviors and as a physical barrier to feeding.