Does R gene resistance allow wheat to prevent plant growth effects associated with Hessian fly (Diptera: Cecidomyiidae) attack?

Resistance genes (R genes) are an important part of the plant's immune system. Among insects, the Hessian fly, Mayetiola destructor (Say) (Diptera: Cecidomyiidae), larva is the target of the greatest number of characterized R genes (H1-H32). The biochemical/molecular mechanism of R gene resistance to Hessian fly is not well understood. In the absence of an effective R gene, larvae caused extensive growth deficits (> 30 cm) in wheat seedlings. In the presence of one of three effective R genes, H6, H9, or H13, larvae caused small growth deficits (approximately 3-4 cm) in two leaves (third and fourth) that were actively growing during the first days of larval attack. After larvae died on R gene plants, the fifth leaf and tiller leaves exhibited small increases in growth (2-4 cm). Growth responses of susceptible and resistant plants diverged at a time when Hessian fly larvae were establishing a nutritive gall tissue at feeding sites. The results of this study support the hypothesis that R gene resistance cannot prevent initial larval attack, but, by stopping the formation of the larval gall, it prevents the most serious consequences of larval attack.     

Resistance of kale varieties to attack by Mamestra brassicae

1 The objectives of this work were to study the resistance of six kale ( Brassica oleracea acephala group) varieties to cabbage moth Mamestra brassicae (L.) expressed as antibiosis and to determine the effect of plant age on larval survival and development.
2 The influence of plant age on resistance was determined using leaves from seedlings and from mature plants. Survival and development of M. brassicae larvae and feeding rates were determined in laboratory bioassays.
3 Leaves from seedlings were more suitable than those of mature plants for establishing differences in resistance. There were significant differences between kale varieties in larval survival, growth rate, leaf feeding, and time to pupation but not pupal weight. The varieties MBG-BRS0031, MBG-BRS0351, and MBG-BRS0287 reduced survival of M. brassicae larvae. Larvae that fed on MBG-BRS0060 were the heaviest and took the longest time to pupation. MBG-BRS0031 was consumed significantly less by larvae than were all the other varieties examined. Leaves from mature plants of MBG-BRS0142 and MBG-BRS0170 were defoliated significantly less than those of other varieties.
4 In conclusion, the variety MBG-BRS0031 may be a promising source of resistance to M. brassicae . Leaf antibiotic resistance was shown to play a role in defense against M. brassicae attack but it is not the only possible mechanism of resistance.     

Avoidance of occupied hosts by the Hessian fly: oviposition behaviour and consequences for larval survival

Abstract 1. Unoccupied wheat plants and wheat plants occupied by conspecific eggs or larvae were presented to ovipositing female Hessian flies in choice tests.
2. The presence of conspecific eggs on the leaf surfaces of wheat plants did not appear to have any effect on the responses of ovipositing Hessian fly females.
3. The presence of conspecific larvae at the base and nodes of wheat plants for 1, 6, or 11 days had significant effects on Hessian fly oviposition. Eggs oviposited on plants were inversely proportional to larval densities and days of larval occupation.
4. Feeding by Hessian fly larvae is associated with several changes in wheat plants. One of these changes, the growth arrestment of the plant, was measured by recording the heights of plants used in oviposition tests. Plant heights were inversely proportional to both larval densities and days of occupation. Plant heights were directly proportional to eggs oviposited on plants.
5. The consequences of adult female avoidance of plants occupied by conspecific larvae were investigated by allowing females to oviposit on unoccupied plants and 1-day, 6-day, and 11-day larval occupied plants, then scoring at the end of the first larval instar the survival of the offspring that resulted from this oviposition.
6. Survival during the first larval instar was 88% for the offspring of females that oviposited on unoccupied plants, decreasing to 82, 31, and 4% on the 1-day, 6-day, and 11-day occupied plant treatments. On these four plant treatments, a positive correlation was found between larval performance (i.e. survival) and the preferences of ovipositing females.
7. On the four plant treatments, relationships between first-instar larval density and first-instar larval survival varied significantly. On unoccupied plants, survival was inversely proportional to density. On plants oviposited on at 6 days of larval occupation, survival was directly proportional to density.     

The influence of greenhouse chrysanthemum on the interaction between the beet armyworm, Spodoptera exigua, and the baculovirus SeMNPV: parameter quantification for a process-based simulation model

During the building of a process-based simulation model for the epidemiology of the multicapsid nucleopolyhedrovirus of S. exigua (SeMNPV) in populations of Spodoptera exigua (Hübner) in greenhouse chrysanthemum, it was found that the effect of host plants had been under-rated. 'Missing links' included (i) the 'natural' background mortality of larvae of S. exigua in practical cropping conditions; (ii) the developmental rate of larvae of S. exigua on plant substrate in a glasshouse as compared to artificial medium in the laboratory; (iii) the validity of the results of dose-mortality and time-mortality bioassays conducted on artificial medium as compared to natural plant substrate; (iv) the distribution of inoculum released from deceased caterpillars over chrysanthemum leaves; and (v) the leaf visit rate of healthy caterpillars (as it affects horizontal transmission). Experiments were carried out to quantify these processes. Developmental rates of S. exigua larvae on greenhouse chrysanthemum were 36% lower than on an artificial diet. The fraction survival during the first, second, third and fourth instar S. exigua larvae in greenhouse chrysanthemum was 0.60, 0.80, 0.88 and 0.95, respectively. Forty percent of the first instar larvae reached the fifth larval stage. Second instar S. exigua larvae reared on chrysanthemum were significantly more susceptible to SeMNPV than larvae reared on an artificial diet. The food source had no effect on the time to kill S. exigua larvae. Cadavers of second, third and fourth instar S. exigua larvae contaminated on average 1.4, 2.5 and 3.3 chrysanthemum leaves. Second to fourth instar S. exigua larvae visited 2–3 leaves per day and spent 15–55% of the time on the underside of leaves. The above information is of critical importance for a trustworthy simulation of the epidemiology of SeMNPV in chrysanthemum.     

No fitness cost for wheat's H gene-mediated resistance to Hessian fly (Diptera: Cecidomyiidae)

Resistance (R) genes have a proven record for protecting plants against biotic stress. A problem is parasite adaptation via Avirulence (Avr) mutations, which allows the parasite to colonize the R gene plant. Scientists hope to make R genes more durable by stacking them in a single cultivar. However, stacking assumes that R gene-mediated resistance has no fitness cost for the plant. We tested this assumption for wheat's resistance to Hessian fly, Mayetiola destructor (Say) (Diptera: Cecidomyiidae). Our study included ten plant fitness measures and four wheat genotypes, one susceptible, and three expressing either the H6, H9, or H13 resistance gene. Because R gene-mediated resistance has two components, we measured two types of costs: the cost of the constitutively-expressed H gene, which functions in plant surveillance, and the cost of the downstream induced responses, which were triggered by Hessian fly larvae rather than a chemical elicitor. For the constitutively expressed Hgene, some measures indicated costs, but a greater number of measures indicated benefits of simply expressing the H gene. For the induced resistance, instead of costs, resistant plants showed benefits of being attacked. Resistant plants were more likely to survive attack than susceptible plants, and surviving resistant plants produced higher yield and quality. We discuss why resistance to the Hessian fly has little or no cost and propose that tolerance is important, with compensatory growth occurring after H gene-mediated resistance kills the larva. We end with a caution: Given that plants were given good growing conditions, fitness costs may be found under conditions of greater biotic or abiotic stress.     

Use of grafting to prevent Hypsipyla grandella (Zeller) (Lepidoptera: Pyralidae) damage to new world Meliaceae species

The susceptible species Cedrela odorata and Swietenia macrophylla to attack by Hypsipyla grandella (Zeller) larvae were grafted onto the resistant species Khaya senegalensis and Toona ciliata. Six-month-old grafted plants were then compared to their reciprocal grafts and to both intact (non-grafted) and autografted plants for damage due to H. grandella larvae and for their effects on larval performance. Two experiments were conducted: one in which the apical bud of the main plant shoot was inoculated with H. grandella eggs, and the other in which the bud was inoculated with third instars. Damage in each experiment was assessed by the number of frass piles, number and length of tunnels, number of damaged leaves, and damage to the apical bud. Larval performance was evaluated in terms of time to reach pupation and pupal weight and length. In both experiments, plant damage differed significantly among treatments (P < 0.03). Resistant rootstocks conferred resistance to susceptible scions. In both experiments, grafting by itself, regardless of the rootstock and scion combination, also reduced damage caused by H. grandella larvae. Scions of autografted susceptible species had similar resistance to susceptible scions grafted on resistant rootstocks. Few larvae reached pupation, and their pupal weight and length were similar.     

Mechanism of resistance to the spotted stalk borer,Chilo sacchariphagus,in the sugarcane cultivar R570

Resistance in sugarcane [Saccharum spec. (Poaceae)] to the spotted stalk borer, Chilo sacchariphagus (Bojer) (Lepidoptera: Pyralidae), was studied by comparing feeding behaviour on resistant cv. R570 and susceptible cv. R579. In a field survey, the feeding behaviour of C. sacchariphagus larvae was described to identify their feeding sites on the plant. In a greenhouse artificial infestation study, we compared the establishment of larvae on potted plants. In laboratory choice and no‐choice experiments, we studied the establishment of larvae on plant organs (stalk, sheath, leaf spindle). Study of the feeding behaviour showed that: (1) first to fourth instars are able to feed on stalk, sheath, and leaf spindle, (2) boring into the stalk occurs mostly in the four uppermost internodes, and (3) most young larvae bore through the abaxial surface of leaf sheaths to reach the stalk. In greenhouse experiments, we observed an early two‐fold reduction of the number of larvae on R570 plants within the first 48 h after infestation. In laboratory experiments, larval antixenosis was demonstrated at the abaxial surface of R570 leaf sheath, but was observed neither in the leaf spindle nor in the stalk. First, second, and third instars were susceptible to this antixenosis. We hypothesize that the main resistance mechanism in R570 is an early reduction of larval establishment on plants, due to antixenosis located at the abaxial surface of leaf sheaths.     

Comparative toxicity of some synthetic insecticides,and a growth inhibitor,in the migratory locust,Locusta migratoria

1. The comparative toxicity of four synthetic insecticides (Dieldrin, Methomyl, Fenitrothion and Permethrin) in the different developmental stages of the migratory locust shows that resistance tends to increase with the developmental larval stages except in the adults which show lower resistance than the fifth larval stage.2. The resistance to a growth inhibitor, Teflubenzuron in the different developmental stages shows the reverse phenomenon—the younger larvae were more resistant than the fourth and fifth larval stages while the adults were almost not affected by this product.3. A comparison of the toxicity of the synthetic insecticides in the two forms of the insect, the solitarious and the gregarious, shows that the solitary insects were more resistant than the gregarious ones.     

Ultrastructural changes in the midguts of Hessian fly larvae feeding on resistant wheat

The focus of the present study was to compare ultrastructure in the midguts of larvae of the Hessian fly, Mayetiola destructor (Say), under different feeding regimens. Larvae were either fed on Hessian fly-resistant or -susceptible wheat, and each group was compared to starved larvae. Within 3 h of larval Hessian fly feeding on resistant wheat, midgut microvilli were disrupted, and after 6 h, microvilli were absent. The disruption in microvilli in larvae feeding on resistant wheat were similar to those reported for midgut microvilli of European corn borer, Ostrinia nubilasis (Hubner), larvae fed a diet containing wheat germ agglutinin. Results from the present ultrastructural study, coupled with previous studies documenting expression of genes encoding lectin and lectin-like proteins is rapidly up-regulated in resistant wheat to larval Hessian fly, are indications that the midgut is a target of plant resistance compounds. In addition, the midgut of the larval Hessian fly is apparently unique among other dipterans in that no peritrophic membrane was observed. Ultrastructural changes in the midgut are discussed from the prospective of their potential affects on the gut physiology of Hessian fly larvae and the mechanism of antibiosis in the resistance of wheat to Hessian fly attack.     

Contribution of fungal biomass to the growth of the shredder, Pycnopsyche gentilis (Trichoptera: Limnephilidae)

1.  1. It has been accepted that aquatic hyphomycetes colonising submerged leaves increase the nutritional value of leaf detritus and suggested that fungal biomass plays a greater role in the growth of shredders than leaf tissue itself. However, it is not clear what proportion of the nutritional needs of shredders is met by fungal biomass.
2.  We fed Pycnopsyche gentilis larvae with tulip poplar ( Liriodendron tulipifera ) leaf discs colonised by the aquatic hyphomycete, Anguillospora filiformis , which had been radiolabelled to quantify the contribution of fungal carbon to the growth of the shredder at different larval developmental stages. Instantaneous growth rates of larvae on this diet were also estimated.
3.  When provided with fungal-colonised leaves (14–16% fungal biomass), the third and the fifth instar larvae of P. gentilis grew at the rates of 0.061 and 0.034 day⁻¹, respectively, but on a diet of sterile leaves, both larval instars lost weight. The incorporation rates of fungal carbon were 31.6 μg C mg⁻¹ AFDM day⁻¹, accounting for 100% of the daily growth rate of the third instar larvae and 8.6 μg C mg⁻¹ AFDM day⁻¹, accounting for 50% of the daily growth rate of the fifth instar larvae.
4.  These results suggest that leaf material colonised by A. filiformis is a high quality food resource for P. gentilis larvae, and that fungal biomass can contribute significantly to the growth of these larvae. Differences in feeding behaviour and digestive physiology may explain the significantly greater assimilation of fungal biomass by the earlier instar than the final instar. To satisfy their nutritional needs the fifth instar larvae would have to assimilate detrital mass that may have been modified by fungal exoenzymes.     

Relative resistance of transgenic tomato tissues expressing high levels of tobacco anionic peroxidase to different insect species.

Different parts of genetically transformed tomato (Lycopersicon esculentum L.) plants that express the tobacco anionic peroxidase were compared for insect resistance with corresponding wild type plants. Leaf feeding by first instar Helicoverpa zea and Manduca sexta was often significantly reduced on intact transgenic plants and/or leaf disks compared to wild type plants, but the effect could depend on leaf age. Leaves of transgenic plants were generally as susceptible to feeding damage by third instar Helicoverpa zea (Boddie) and Manduca sexta (L.) as wild type plants. Green fruit was equally susceptible to third instar larvae of H. zea in both type plants, but fruit of transgenic plants were more resistant to first instar larvae as indicated by significantly greater mortality. Basal stem sections were more resistant to neonate larvae of H. zea and adults of Carpophilus lugubris Murray compared to wild type plants as indicated by significantly greater mortality and/or reduced feeding damage. Thus, tobacco anionic peroxidase activity can increase plant resistance to insects in tomato, a plant species closely related to the original source plant species, when expressed at sufficiently high levels. However, the degree of resistance is dependent on the size of insect and plant tissue involved.     

不同光照强度下兴安落叶松对舞毒蛾幼虫生长发育及防御酶的影响

为了探讨舞毒蛾对光照环境变化引起的落叶松抗性的变化是如何响应的,分析了不同光照强度处理的兴安落叶松对舞毒蛾幼虫生长发育状况和生理指标变化的影响。研究发现,取食50%和25%光照强度下生长的落叶松后,舞毒蛾幼虫平均体重、蛹重、化蛹率和羽化率与对照组相比均显著下降（P<0.05）。幼虫体内保护酶SOD、POD、CAT和解毒酶ACP、AKP、CarE、GSTs、MFO活性与对照组(100% 光照)相比均显著降低（P<0.05）且除CAT外在50%光照强度下降低最为显著（P<0.05）;4龄和5龄幼虫取食同一光强处理的落叶松后,SOD、POD、CarE、GSTs和MFO活性5龄显著低于4龄（P<0.05）,ACP和AKP活性5龄却显著高于4龄（P<0.05）,表明昆虫在不同的发育阶段启用不同的排毒酶系。光照差异对取食人工饲料的舞毒蛾生长发育和排毒代谢酶活性均没有显著影响,说明舞毒蛾生长发育和酶活性的变化是因寄主植物抗性的影响而非光照强度的作用。本研究结果表明,采取适当的营林措施调节林分内的光照条件,可以提高落叶松的自主抗虫性,增强其抵御害虫危害的能力,有效控制害虫危害,减少化学农药的施用量。     

Hessian fly Avirulence gene loss‐of‐function defeats plant resistance without compromising the larva's ability to induce a gall tissue

Plant pathogen effectors encoded by Avirulence (Avr) genes benefit the pathogen by promoting colonization and benefit plants that have a matching resistance (R) gene by constituting a signal that triggers resistance. The Hessian fly, Mayetiola destructor (Say) (Diptera: Cecidomyiidae), resembles a plant pathogen in showing R/Avr interactions. Because of these interactions, a wheat plant with the H13 resistance gene can be resistant or susceptible depending on the genotype of the larva that attacks the plant, being resistant if attack comes from a larva with a functional vH13 gene, but susceptible if attack comes from a larva with a non‐functional vH13 gene. In this study we asked: does this susceptible interaction involving plants with H13 look like susceptible interactions with plants lacking H13? Possibly, the H13 plant attacked by a larva with a non‐functional vH13 is induced to partial rather than complete resistance. Or the larva, lacking its vH13‐encoded effector, is compromised in its ability to induce susceptibility, which includes forcing the plant to create a gall nutritive tissue. Responses of epidermal cells to larval attack were explored using imaging techniques (light microscopy, scanning and transmission electron microscopy). Whole‐organism responses were investigated by measuring the growth of plants and larvae. No evidence was found for partial resistance responses by H13 plants or for a compromise in the ability of vH13 loss‐of‐function larvae to induce susceptibility. It appears that disrupting vH13 function is sufficient for overcoming the induced resistance mediated by the H13 gene.     

A novel wheat gene encoding a putative chitin-binding lectin is associated with resistance against Hessian fly

The gene-for-gene interaction triggering resistance of wheat against first-instar Hessian fly larvae utilizes specialized defence response genes not previously identified in other interactions with pests or pathogens. We characterized the expression of Hfr-3 , a novel gene encoding a lectin-like protein with 68–70% identity to the wheat germ agglutinins. Within each of the four predicted chitin-binding hevein domains, the HFR-3 translated protein sequence contained five conserved saccharide-binding amino acids. Quantification of Hfr-3 mRNA levels confirmed a rapid response and gradual increase, up to 3000-fold above the uninfested control in the incompatible interaction 3 days after egg hatch. Hfr-3 mRNA abundance was influenced by the number of larvae per plant, suggesting that resistance is localized rather than systemic. In addition, Hfr-3 was responsive to another sucking insect, the bird cherry-oat aphid, but not to fall armyworm attack, wounding or exogenous application of methyl jasmonate, salicylic acid or abscisic acid. Western blot analysis demonstrated that HFR-3 protein increased in parallel to mRNA levels in crown tissues during incompatible interactions. HFR-3 protein was detected in both virulent and avirulent larvae, indicating ingestion. Anti-nutritional proteins, such as lectins, may be responsible for the apparent starvation of avirulent first-instar Hessian fly larvae during the initial few days of incompatible interactions with resistant wheat plants.     

Influence of plant silicon and sugarcane cultivar on mandibular wear in the stalk borer Eldana saccharina

1 Silicon can increase the resistance of plants to attack by herbivorous insects. The present study aimed to determine the effect of silicon and cultivar on mandibular wear in larvae of the sugarcane stalk borer Eldana saccharina Walker (Lepidoptera: Pyralidae).
2 Four sugarcane cultivars, resistant (N21, N33) and susceptible (N11, N26) to E. saccharina were grown in a pot trial in silicon deficient river sand, with (Si+) and without (Si−) calcium silicate. Individual third-instar larvae were confined on the sugarcane stalk at three known feeding sites (leaf bud, root band and internode) and left to feed for 21 days.
3  Eldana saccharina larval heads were mounted on stubs, with the mandibles oriented horizontally and photographed under a scanning electron microscope. Mandibular wear was measured from the digital images using a quantitative method.
4 Although there was a trend for increased wear in larvae that developed on Si+ cane, no significant effect of silicon, cultivar or site on mandibular wear of E. saccharina was shown.
5 This is the first study to accurately and quantitatively measure the mandibular wear of an insect fed on Si+ plants.