Wheat bulb fly,Delia coarctata,larval attraction to phenolic components of host‐plant root exudates

Wheat bulb fly (WBF), Delia coarctata Fallén (Diptera: Anthomyiidae), larvae are a subterranean pest of wheat [Triticum aestivum L. (Poaceae)] and other cereals. Larvae locate host plants through chemotaxis and chemokinesis, utilising the primary plant metabolite carbon dioxide as a ‘search trigger’ and Poaceae‐specific secondary plant metabolites exuded from the plant. The aim of this study was to use arena bioassays to identify further compounds involved in the host‐finding process. The larval behavioural response to four concentrations of syringic and vanillic acid, chemical constituents of host‐plant exudates, were tested. Analysis of the final resting position of D. coarctata larvae by the Rayleigh test of uniformity identified attraction to wheat seedling exudates and to both compounds at the lowest concentrations tested, with syringic acid concentrations being most attractive at 0.1 mg l^?1 and vanillic acid being most attractive at 0.001 mg l^?1. These results add more detail to the subterranean chemical ecology of this species, allowing a behavioural sequence for host‐plant orientation by WBF larvae to be proposed.     

Searching for wheat resistance to aphids and wheat bulb fly in the historical Watkins and Gediflux wheat collections

Insect pests can reduce wheat yield by direct feeding and transmission of plant viruses. Here we report results from laboratory and field phenotyping studies on a wide range of wheat, including landraces from the Watkins collection deriving from before the green revolution, more modern cultivars from the Gediflux collection (north‐western Europe) and modern UK Elite varieties, for resistance to the bird cherry‐oat aphid, Rhopalosiphum padi (Homoptera: Aphididae) and the English grain aphid, Sitobion avenae (Homoptera: Aphididae). A total of 338 lines were screened for R. padi and 340 lines for S. avenae. Field trials were also conducted on 122 Watkins lines to identify wheat bulb fly, Delia coarctata, preference on these landraces. Considerable variation was shown in insect performance among and within different wheat collections, with reduced susceptibility in a number of varieties, but phenotyping did not identify strong resistance to aphids or wheat bulb fly. Field trials showed within collection differences in aphid performance, with fewer aphids populating lines from the Watkins collection. This differs from development data in laboratory bioassays and suggests that there is a pre‐alighting cue deterring aphid settlement and demonstrates differences in aphid preference and performance on older plants in the field compared with seedlings in the laboratory, highlighting the need for phenotyping for aphid resistance at different plant growth stages. No association was identified between performance of the different insect species on individual varieties, potentially suggesting different nutritional requirements or resistance mechanisms.     

Effects of activated charcoal on attack by larvae of the wheat bulb fly, Delia coarctata

Activated charcoal interferes with normal host-seeking by larvae of the wheat bulb fly, Delia coarctata, by adsorbing arrestant compounds exuded from the plants. Thus, in laboratory tests, activated charcoal in soil surrounding one of two rows of wheat seedlings decreased larval attack on the treated row. Charcoal in soil above the level of the seeds decreased attack but had no effect when below the seeds. However, there was no decrease in attack when both rows were in charcoal. Attempts to develop a practical application of this effect in the field are described.     

Intercropping impacts the host location behaviour and population growth of aphids

Increasing intrafield plant diversity has been shown to regulate pest populations in various agroecosystems. Among the suggested mechanisms for this bottom-up pest control, the disruptive crop hypothesis states that herbivores' abilities to locate and colonize their host plants are reduced by the presence of non-host plants. Under laboratory conditions, we evaluated how intercropping wheat and legumes modifies the behaviour of apterous cereal aphids, Sitobion avenae (Fabricius) (Hemiptera: Aphididae), in terms of host plant location and population growth. We compared two intercropping systems – soft winter wheat, Triticum aestivum L. (Poaceae), associated with winter pea, Pisum sativum L., or with white clover, Trifolium repens L. (both Fabaceae) – and sole stands of soft winter wheat. Aphids needed more time to locate their wheat host plant and then spent less time on wheat when it was intercropped with clover. At the population level, and accounting for host plant biomass, only intercropping wheat with clover significantly reduced aphid densities on wheat, as this was particularly disruptive to S. avenae behaviour and population growth. Our laboratory study points out that the species used as non-host plants and their density are important parameters that should be taken into account in field studies on intercropping systems.     

Development of a pest threshold decision support system for minimising damage to winter wheat from wheat bulb fly,Delia coarctata

Wheat bulb fly, Delia coarctata, is an important pest of winter wheat in the UK, causing significant damage of up to 4 t/ha. Accepted population thresholds for D. coarctata are 250 eggs/m² for crops sown up to the end of October and 100 eggs/m² for crops sown from November. Fields with populations of D. coarctata that exceed the thresholds are at higher risk of experiencing economically damaging infestations. In the UK, recent withdrawal of insecticides means that only a seed treatment (Signal 300 ES) is available for chemical control of D. coarctata; however, this is only effective for late-sown crops and accurate estimations of annual population levels are required to ensure a seed treatment is applied if needed. As a result of the lack of postdrilling control strategies, the management of D. coarctata is becoming reliant on nonchemical methods of control. Control strategies that are effective in managing similar stem-boring pests of wheat include sowing earlier and using higher seed rates to produce crops with greater pest tolerance. In this study, we develop two predictive models that can be used for integrated D. coarctata management. The first is an updated pest level prediction model that predicts D. coarctata populations from meteorological parameters with a predictive accuracy of 70%, a significant improvement on previous prediction models. Our second model predicts the maximum number of shoots for a winter wheat crop that would be expected at the terminal spikelet development stage. This shoot number model uses information about the thermal time from plant emergence to terminal spikelet, leaf phyllochron length, plant population and sowing date to predict the degree of tolerance a crop will have against D. coarctata. The shoot number model was calibrated against data collected from five field experiments and tested against data from four experiments. Model testing demonstrated that the shoot number model has a predictive accuracy of 65.7%. The foundation for a future decision support system using these models for the sustainable management of D. coarcata risk is described. It should be noted that these models represent a stepping-stone towards a decision support system and that further model validation over a wider geographic range is required.     

A comparison of a corer and a shovel for sampling populations of wheat bulb fly Delia coarctata eggs

The most commonly used tool for sampling soil for eggs of the wheat bulb fly Delia coarctata Fall.) is a corer, but in Scotland samples are taken with a modified shovel. The sampling properties of a corer and shovel were compared in two exercises in which soil samples were taken from the ridge, sides and furrow of drills of maincrop potatoes at several sites in east Scotland. Although each tool sampled approximately the same surface area of soil, the corer sampled three times as much soil as the shovel. The numbers of wheat bulb fly eggs estimated by the corer and shovel were similar in both sampling exercises and there were no indications that the comparison of the two tools was affected by the sampling position on the potato ridge. The results of the two exercises gave no reason to believe that in similar conditions the shovel is an inferior sampling tool or that population estimates obtained with a shovel in Scotland are not comparable with those obtained with a corer. Evidence from some sites suggested that wheat bulb flies lay more eggs on the potato ridge than in the furrow.     

A comparison of wheat bulb fly (Delia coarctata) egg numbers in ridge and furrow soil samples from potato crops

In soil samples taken with a shovel from 97 potato maincrops in east Scotland during 1978 to 1981, wheat bulb fly (Delia coarctata) eggs were more numerous on ridges than in furrows. The ratio of the number of eggs on the ridge to the number of eggs in the furrow was higher in Lothian Region (3·25) than in Fife Region (1·59) but differed significantly (P < 0·05) from 1·0 in both Regions. To obtain maximum precision of the estimate of the total wheat bulb fly egg population in a potato field, more samples should be taken from the ridge than from the furrow, in approximate proportion to the egg numbers.     

The invasion and development of the cereal cyst-nematode, Heterodera avenae in the roots of autumn-and spring-sown cereals

Observations were made at 2 or 4 wk intervals from December to harvest on all stages of Heterodera avenae in winter oats growing on infested land. Second-stage larvae were present in all soil samples except on 5 and 20 July. Invasion and development of larvae was slow during winter. The nodal and seminal roots of winter oats were both heavily invaded by the nematode; larvae which invaded seminal roots tended to become male whereas those in nodal roots tended to become female. There was a small second invasion in August. Females were first observed on the roots of winter oats on 17 May, 214 days after the crop was sown and 62 days after the first fourth-stage larva was observed. The nodal roots of spring barley contained few H. avenae larvae whereas these roots were heavily invaded in winter wheat and oats. In spring barley the nodal roots were developing in June and July when few second-stage larvae were in the soil whereas in winter oats and wheat the nodal roots were growing rapidly in April when larvae were most numerous, and so were heavily invaded.     

EXPERIMENTS ON THE MODE OF ACTION OF DIELDRIN SEED-DRESSINGS AGAINST SEEDLING ATTACK BY THE LARVAL FRIT FLY, OSCINELLA FRIT L

Experiments with dieldrin seed-dressings were done to see whether the conditions favouring their action against wheat bulb fly larvae apply also to frit fly.
Experiments with oats and wheat in boxes and a small field trial with oats showed that fewer newly hatched larvae were killed when dieldrin-treated seed was sown deeply than when sown 1/4 - 1/2 in. deep; shallow sowing probably puts the insecticide where the larvae are likely to meet it before they attack the plant. Kill was never complete even with dieldrin applied at almost forty times more than the highest rate normally used. This is partly because some larvae hatch from eggs on the plant and these, unlike those from soil eggs, can enter the shoot without touching the seed dressing.
Even in the most favourable circumstances (large doses of dieldrin applied to the shallow-sown seed and the plant at the one-leaf stage at the time of attack) there was little evidence that frit fly larvae are killed inside the shoot by systemic action of dieldrin translocated from the seed. This is not because the pales which normally surround the oat seed impair the uptake of insecticide, or because of any other factor specific to oats. Frit fly larvae also survived in wheat from seed dressed with doses of dieldrin that can kill almost all wheat bulb fly larvae by systemic action. Possible reasons for this difference are discussed.     

Chemistry of components in wheat and oats that influence behaviour of wheat bulb fly larvae

Chemicals produced by wheat and oats influence the behaviour of wheat bulb fly larvae; in laboratory tests, wheat extracts ‘arrest’ these larvae whereas extracts from oats (a non-host plant) have an ‘anti-arrestant’ effect. The active compounds from both types of extract have been purified and their chemistry investigated. The results suggest that the wheat ‘arrestant’ is a polyphenolic compound, possibly as a glycoside, and the oat ‘anti-arrestant’ a polyhydroxylated aliphatic material.     

II. Cereal disease: the interplay between resistance and pathogenicity: Inheritance of pathogenicity in Gaeumannomyces graminis var. tritici

Variation in host response of isolates of the eyespot pathogen from different sources was examined over a number of years. Pathogen types were found in intensively-cropped couch-infested cereal sites that were almost as virulent on Agropyron repens (couch) as on wheat or barley. The commonly occurring wheat (W) type isolates from couch-free cereal crops were virulent on wheat and barley but avirulent on couch. Couch (C) types were isolated not only from couch but also from wheat, barley and oat crops with couch infestation. In pathogenicity tests on rye, C. types did not differ in virulence from the more commonly occurring W types. Aegilops ventricosa was equally resistant to both types. W type isolates from wheat and barley were examined to assess differential pathogenicity on wheat and barley. Sequential cropping with single cereal crops was used to separate out possible specific types. Isolates from fourth wheat and fourth barley crops were more pathogenic on the original than on the alternative host. When comparisons were made between isolates from third and fifth consecutive wheat and barley crops only those from barley showed a preference for the original host. An experiment comparing isolates from third and seventh consecutive wheat and barley crops showed a decline in virulence from the short to the longer sequences on the alternative but not on the original host.     

Feeding by Hessian Fly (Mayetiola destructor [Say]) Larvae on Wheat Increases Levels of Fatty Acids and Indole-3-Acetic Acid but not Hormones Involved in Plant-Defense Signaling

Gall-inducing insects exert a unique level of control over the physiology of their host plants. This control can extend to host–plant defenses so that some, if not most, gall-inducing species appear to avoid or modify host plant defenses to effect production of their gall. Included among gall insects is Hessian fly (Mayetiola destructor [Say], Diptera: Cecidomyiidae), a damaging pest of wheat (Triticum aestivum L.) and an emerging model system for studying plant–insect interactions. We studied the dynamics of some defense-related phytohormones and associated fatty acids during feeding of first instar Hessian fly larvae on a susceptible variety of wheat. We found that Hessian fly larvae significantly elevated in their host plants’ levels of linolenic and linoleic acids, fatty acids that may be nutritionally beneficial. Hessian fly larvae also elevated levels of indole-3-acetic acid (IAA), a phytohormone hypothesized to be involved in gall formation, but not the defense-related hormones jasmonic (JA) and salicylic acids. Moreover, we detected in Hessian fly-infested plants a significant negative relationship between IAA and JA that was not present in control plants. Our results suggest that Hessian fly larvae may induce nutritionally beneficial changes while concomitantly altering phytohormone levels, possibly to facilitate plant-defense avoidance.     

The interaction of a Botanophila fly species with an exotic Epichloë fungus in a cultivated grass: fungivore or mutualist?

Epichloë spp. (Ascomycetes: Clavicipitaceae) are endophytic fungii of Pooid grasses that cause choke disease, the suppression of seed production. They also host Botanophila spp. (Diptera: Anthomyiidae), the larvae of which feed on the fungus. Studies on Epichloë elymi on wild grasses indicate that the flies transfer spermatia between Epichloë mating types, thereby affecting cross‐fertilization, suggesting that the fungus–fly interaction reflects obligatory mutualism. Epichloë typhina, inadvertently introduced into western USA, was first detected in cultivated Dactylis glomerata L. fields in 1996. It spread rapidly, raising concerns about impacts on seed production. The present study was conducted to address questions pertaining to the occurrence and nature of the fungus–fly interaction in the new habitat of E. typhina. The first report of an endemic Botanophila species associated with E. typhina in Oregon is presented here. Surveys of D. glomerata fields indicated no correlation between fly abundance and fungal fertilization. In one field, no fly eggs or larvae were detected, but fertilized stromata were universally present. The fly was established in the remaining 12 fields surveyed, but while the number of stromata with fly larvae ranged from 6 to 98%, stromata development was uniform. Up to 10 larvae were present on a stroma, and these consumed >90% of the perithecia. Comparisons of pupal weights indicated that the fungal resource was not limiting, even at high larval densities. An exclusion study in a D. glomerata field also indicated that E. typhina fertilization occurred without the fly. In Oregon, the fly clearly benefits from the association with the fungus, but there is no evidence of benefit to the fungus. Thus if obligatory mutualism in the fungus–fly relationship described from the midwestern USA is the norm, our studies suggest a shift in the interaction to one of simple foraging on the fungus by fly larvae.     

Companion planting – do aromatic plants disrupt host‐plant finding by the cabbage root fly and the onion fly more effectively than non‐aromatic plants?

Brassica and Allium host‐plants were each surrounded by four non‐host plants to determine how background plants affected host‐plant finding by the cabbage root fly (Delia radicum L.) and the onion fly [Delia antiqua (Meig.)] (Diptera: Anthomyiidae), respectively. The 24 non‐host plants tested in field‐cage experiments included garden ‘bedding’ plants, weeds, aromatic plants, companion plants, and one vegetable plant. Of the 20 non‐host plants that disrupted host‐plant finding by the cabbage root fly, fewest eggs (18% of check total) were laid on host plants surrounded by the weed Chenopodium album L., and most (64% of check total) on those surrounded by the weed Fumaria officinalis L. Of the 15 plants that disrupted host‐plant finding in the preliminary tests involving the onion fly, the most disruptive (8% of check total) was a green‐leaved variant of the bedding plant Pelargonium × hortorum L.H. Bail and the least disruptive (57% of check total) was the aromatic plant Mentha piperita × citrata (Ehrh.) Briq. Plant cultivars of Dahlia variabilis (Willd.) Desf. and Pelargonium×hortorum, selected for their reddish foliage, were less disruptive than comparable cultivars with green foliage. The only surrounding plants that did not disrupt oviposition by the cabbage root fly were the low‐growing scrambling plant Sallopia convolvulus L., the grey‐foliage plant Cineraria maritima L., and two plants, Lobularia maritima (L.) Desv. and Lobelia erinus L. which, from their profuse covering of small flowers, appeared to be white and blue, respectively. The leaf on which the fly landed had a considerable effect on subsequent behaviour. Flies that landed on a host plant searched the leaf surface in an excited manner, whereas those that landed on a non‐host plant remained more or less motionless. Before taking off again, the flies stayed 2–5 times as long on the leaf of a non‐host plant as on the leaf of a host plant. Host‐plant finding was affected by the size (weight, leaf area, height) of the surrounding non‐host plants. ‘Companion plants’ and aromatic plants were no more disruptive to either species of fly than the other plants tested. Disruption by all plants resulted from their green leaves, and not from their odours and/or tastes.     

Change in response of Rhopalosiphum padi spring migrants to the repellent winter host component methyl salicylate

Olfactometry showed that the response of spring migrants of the bird cherry-oat aphid, Rhopalosiphum padi (L.) (Homoptera: Aphididae), to the repellent winter host volatile methyl salicylate changes with age of the adult aphid. Between three and four days after becoming adult, and having left the winter host Prunus padus L., aphids lost their negative response to the chemical. The change in response was not associated with contact with a summer host, oats. In a settling choice bioassay, migrants avoided oats which had been exposed to volatile methyl salicylate. Aphids with removed antennal tips did not avoid the exposed plant, indicating that plant choice was influenced by cues from the plant surface. The results are discussed in relation to the use of methyl salicylate in integrated control.     

Olfactory response of three parasitoid species (Hymenoptera: Braconidae) to volatiles of guavas infested or not with fruit fly larvae (Diptera: Tephritidae)

The olfactory responses of the native parasitoids Doryctobracon areolatus (Szépligeti) and Asobara anastrephae (Muesebeck) and of the exotic parasitoid Diachasmimorpha longicaudata (Ashmead) to guava (Psidium guajava L.) infested or not with fruit fly larvae were evaluated. D. areolatus and D. longicaudata females responded to the odors of uninfested rotting guavas, although D. areolatus was also attracted to fruits at the initial maturation (turning) stage. The females of these species recognized the volatiles of guavas containing Ceratitis capitata (Wied.) larvae. However, in bioassays involving fruits with larvae of different instars, D. longicaudata females were not able to separate between fruits containing C. capitata larvae at the initial instars and larvae at the third instar. In the evaluations of volatiles released by guavas containing C. capitata and Anastrepha fraterculus (Wied.) larvae, the D. longicaudata females were oriented toward the volatiles of fruits containing both host species, but differed significantly from volatiles of guavas containing C. capitata larvae. The D. areolatus females also showed responses to both species, although with a preference for volatiles of fruits containing A. fraterculus larvae. The A. anastrephae females were oriented toward the odors of fruits infested with both fruit fly species. In the shade house, D. longicaudata females were oriented to volatiles of rotting fruits containing larvae or not, but could not significantly differentiate between hosts. D. areolatus females were not attracted toward fruits on the ground in the shade house, regardless of host, suggesting that this parasitoid does not forage on fallen fruits.     

Localization of Proteins Immunologically Related to Subunits of Stress 310-kD Protein in Winter Wheat Mitochondria

We studied the localization of polypeptides immunochemically related to subunits of cold-shock 310-kD protein from winter rye (Secale cerealeL.) in mitochondria and submitochondrial structures of winter wheat (Triticum aestivumL.) seedlings. Polypeptides were separated by SDS-PAGE and probed with the antibody against 310-kD protein from rye seedlings. Wheat mitochondria contained the following polypeptides cross-reacting with this antibody: 66, 60, 55, and 23 kD in the inner membrane; 60 and 58 kD in the outer membrane; and 66 and 55 kD in the matrix.     

Fungal volatiles associated with moldy grain in ventilated and non-ventilated bin-stored wheat

The fungal odor compounds 3-methyl-l-butanol, l-octen-3-ol and 3-octanone were monitored in nine experimental bins in Winnipeg, Manitoba containing a hard red spring wheat during the autumn, winter and summer seasons of 1984–85. Quality changes were associated with seed-borne microflora and moisture content in both ventilated and non-ventilated bins containing wheat of 15.6 and 18.2% initial moisture content. All three odor compounds occurred in considerably greater amounts in bulk wheat in non-ventilated than in ventilated bins, particularly in those with wheat having 18.2% moisture content. The presence of these compounds usually coincided with infection of the seeds by the fungi Alternaria alternata (Fr.) Keissler, Aspergillus repens DeBarry, A. versicolor (Vuill.) Tiraboschi, Penicillium crustosum Thom, P. oxalicum Currie and Thom, P. aurantiogriseum Dierckx, and P. citrinum Thom. High production of all three odor compounds in damp wheat stored in non-ventilated bins was associated with heavy fungal infection of the seeds and reduction in seed germinability. High initial moisture content of the harvested grain accelerated the production of all three fungal volatiles in non-ventilated bins.     

Diurnal pattern of death and sporulation in Entomophaga maimaiga-infected Lymantria dispar

Entomophaga maimaiga Humber, Shimazu, et Soper (Zygomycotina: Entomophthoraceae) is a naturally occurring obligate fungal pathogen specific to gypsy moth, Lymantria dispar (L.) (Lepidoptera: Lymantriidae) larvae. This fungus is considered the most important natural enemy of this pest insect in North America and Asia. A critically important step for the development of E. maimaiga epizootics is the transmission of propagules to healthy larvae, a process known to require high humidity. Some pathogens are known to manipulate the time of day that hosts die so that propagules are produced to maximize chances of survival and thus enhance transmission. The objective of this study was to assess whether E. maimaiga manipulates L. dispar to die at a certain time of day. Laboratory bioassays were conducted at 15 and 20 °C to record the 24‐h activity pattern of death and sporulation exhibited under an L14:D10 photoperiod and 100% r.h. by four isolates of E. maimaiga in its host L. dispar. Events were recorded every 4 h. Our results clearly demonstrate that E. maimaiga‐infected L. dispar larvae die mainly in the afternoon and that the fungus sporulates during the night. The rhythm was independent of the fungal isolate tested and type of spores produced after larval death. By raising the temperature from 15 to 20 °C, the peak death time narrowed and sporulation was initiated earlier at night.     

Archives of insect biochemistry and physiology guide for authors

Several carbohydrases and glycosidases from the alimentary cancal and/or salivary glands of feeding larvae of mayetiola destructor have been identified. Pectinase activity was identified in the midgut and may be present in the salivary glands. No endocellulase activity was found in larvae; however, hemicellulase activity was detected in extract of larvae. Amylase activity was present in midguts from feeding larvae and at a low level in extract of salivary glands. Amylases detected in the midgut showed mobilities during polyacrylamide gel electrophoresis similar to the two major amylases in tissues of the insect's host plant. The possibility exists that Hessian fly larvae utilize amylases obtained from their host plant in the digestion of starch. The major glycosidases detected in the midgut lumen of larve were: α-D-glucosidase and α-D-and β-D-galactosidase. The role of these enzymes in the feeding process of Hessian fly larvae is discussed as well as their potential role in feeding damage to wheat.