Oviposition behavior of the cabbage root fly,Delia radicum (L.), influenced by host plant extracts

An ethanolic extract of cabbage leaves (Brassica oleraceavar. capitata,Golden Acre)and derived fractions were tested on natural and surrogate leaves in order to study the oviposition behavior of the cabbage root fly Delia radicum(Diptera: Anthomyiidae). On surrogate plastic leaves coated with a thin layer of paraffin wax and treated with 0.1 gram leaf equivalent (gle) of an ethanolic raw cabbage extract, the females displayed the same sequence of behavioral patterns as on a natural host plant. The quantified oviposition behavior correlated well with the actual number of eggs laid. The extract-treated surrogate leaves received at least as many eggs as natural leaves with a similar surface area. Previous exposure to surrogate or natural leaves seemed not to induce a specific preference. The three fractions (hexane, butanol, and water) of the raw extract stimulated the oviposition behavior. This was taken as evidence for the presence of a multicomponent mixture or a chemical pattern in the cabbage leaves stimulating oviposition. At the tested concentration each fraction alone could stimulate in some individuals the complete behavior. No effect of volatile components of the raw extract could be detected. This is thought to be due to the extraction procedure, which limits the production of attractive, volatile compounds, such as isothiocyanates.     

Foliar form, colour and surface characteristics influence oviposition behaviour in the cabbage root fly Delia radicum

The effects of foliar form, colour and surface characteristics on the oviposition behaviour of the cabbage fly were investigated by presenting females with various leaf models made of paper. The models differed in shape, size, colour, surface coating, the presence of 3 dimensional folds and a stem. It was shown that in the presence of host leaf surface extracts physical factors can strongly influence oviposition behaviour. Females laid the most eggs around the base of bright green or yellow models having a stem, vertical folds and covered with a thin layer of paraffin. The vertical folds in the surface increased the chance of a behavioural transition from leaf run to stem run.

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Zusammenfassung Mit Hilfe von Papiermodellen von Kohlblättern wurde der Einfluss von visuellen und mechanischen Eigenschaften des Blattes auf das Eiablageverhalten der kleinen Kohlfliege, Delia radicum, untersucht. Die verwendeten Modelle unterschieden sich in Form, Grösse, Farbe, Oberflächen struktur und Stiel des Blattes. Es wurde gezeigt, dass physikalische Reize zusammen mit den chemischen Stoffe auf der Blattoberfläche stimulierend wirken. Ein natürliches Eiablageverhalten und die stärkste Eiablage wurde durch ein Modell ausgelöst, welches sich durch leuchtend grüne Farbe, Stiel und vertikale Falten und einen Paraffinnüberzug auszeichnete.

     

The effectiveness of traps used currently for monitoring populations of the cabbage root fly (Delia radicum)

Six different types of traps were compared for capturing adults of Delia radicum. Cone traps caught so few flies that it seems inadvisable for them to be used for monitoring populations of this pest. All the other traps tested were suitable for monitoring D. radicum populations. When expressed on a trap for trap basis, the large 1800 cm² Canadian traps caught most flies. When expressed as the numbers of flies caughthnit area of trapping surface, most flies were caught in the water traps. Similar numbers of D. radicum, D. platura, syrphids, blowflies and a tachinid, Eriothrix rufomaculatus were caught per unit area on each of three different vertical sticky traps. Although water traps have the advantage that they catch about 5 times as many females D. radicum/unit area as the most effective vertical sticky traps of a comparable size, their disadvantage is that they catch about 10 times as many beneficial syrphids. The reasons why water traps are so effective against D. radicum are discussed, with details of how to convert water-trap data to sticky-trap equivalents, and vice versa.     

The olfactory response of cabbage root fly Delia radicum to the host plant volatile allylisothiocyanate

The responses of gravid female cabbage root flies, Delia radicum (L.), to brassica odour and allylisothiocyanate (ANCS) presented in either a discrete plume or uniformly dispersed in air passing through a wind tunnel were investigated. Relatively fast, straight flights occurred in diffuse odour conditions, while in discrete plumes flights were slower with more frequent changes of direction.Flies maintained a constant ground speed and track angle over the straight legs of their flight tracks in two different wind speeds by changing air speed and course. As flies approached an odour source in a discrete plume ground speed was progressively decreased and frequency of turning increased but track angles remained constant. This was achieved by reducing air speed and course angle.Flies released at increasing distances from an odour source moved upwind towards it in a series of short flights, the number relating to distance.The results are discussed in the light of current theories of anemotactic odour source locations and a sequence of orientated behaviour for host plant finding is proposed.

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Les réactions olfactives de Delia radicum à l'allylisothiocyanate volatil émis par les plantes-hôtes

Résumé L'étude a porté sur les réactions de femelles fécondées de Delia radicum L. à l'odour de chou et à l'allylisothiocyanate (ANCS) offerts, soit sous forme d'un fin panache, soit uniformément répartis dans l'air traversant un tunnel à vent.Avec une odeur diffuse les vols étaient relativement rapides et rectilignes, tandis qu'avec un fin panache les vols étaient plus lents avec des changements de direction plus fréquents.En présence de deux vitesses du courant d'air les mouches ont maintenu leur vitesse au sol et leur angle de vol par rapport à la direction du déplacement constants en modifiant leur vitesse de vol et leur direction.Quand les mouches approchaient de la source odorante dans un fin panache, la vitesse au sol diminuait progressivement et la fréquence des virages augmentait, les angles de vol restant constant. Ceci était obtenu en réduisant de la vitesse de l'air et l'angle du déplacement.Les mouches paraissent utiliser une anémotaxie optomotrice et mécanique pendant la localisation de l'hôte. Les résultats sont discutés à la lumière des théories classiques sur la localisation anémotactique des sources odorantes.

     

Host-stage selection by Trybliographa rapae, a parasitoid of the cabbage root fly Delia radicum

Host-stage selection by Trybliographa rapae Westwood (Hymenoptera: Figitidae) was studied in choice and no-choice experiments in the laboratory. The parasitoid was able to reproduce in first, second, and third instars of the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae), but oviposition occurred more frequently in third instars when all three developmental stages were offered simultaneously. Oviposition in third instars increased the rate of development of offspring and their body size, but did not alter sex ratio. Results are discussed in the light of predictions made by the theory of optimal host acceptance.     

Effect of trap background on cabbage root fly landing and capture

Pieces (600 mm × 800 mm) of coloured board, plastic sheeting and woven materials, place beneath water traps prevented the traps from becoming soiled during rainy weather. Such backgrounds are not recommended for use with traps for monitoring populations of the cabbage root fly (Delia radicum L. — Diptera: Anthomyiidae), however, as, instead of increasing trap catch they reduced the numbers of female flies caught by 70%–90%. The main effect was that the visually attractive stimuli from the introduced backgrounds competed with those from the trap. A white background competed with a white trap on a direct fly/unit area basis. Green backgrounds stimulated males to land and the vertical stems of both real and artificial grass induced trivial flights that resulted in greater numbers of males entering traps resting on short grass. Most females were caught over bare soil. To minimize the variation in catch between traps used for monitoring cabbage root fly populations, the background beneath each trap should be similar. For maximum capture, the background should be of grass for male flies and of bare soil for female flies. Care is required if data from traps within mulched crops are used to make pest control decisions, as they will underestimate considerably the numbers of flies in such crops.     

Possible impact of global warming on cabbage root fly (Delia radicum) activity in the UK

A program for simulating the patterns of egg-laying by populations of the cabbage root fly was used to model the effects of global warming on future cabbage root fly attacks. An increase of 3°C in mean daily temperature would cause the cabbage root fly to become active about a month earlier in the year than at present. Under such conditions, the emergence of flies from the overwintering population would be less synchronised, as the completion of diapause and post-diapause development would occur at the same time in different individuals within the population. However, there would continue to be only three generations of fly each year, even in the south of England. With temperature increases of 5°C or 10°C, the fly would complete four generations each year and aestivation would seriously disrupt egg-laying. These rises in temperature would have a major impact on cabbage root fly activity and would require new strategies for controlling this pest.     

An analysis of the deterrent effect of aphids on cabbage root fly (Delia radicum) egg-laying

Abstract. 1. The cabbage root fly, Delia radicum (L.), was deterred from laying eggs on brassica plants with >250 cabbage aphid, Brevicoryne brassicae (L.), or peach-potato aphid, Myzuspersicae (Sulz.).
2. Flies did not lay on plants infested with >250 aphids.
3. Preparations of (E)-β-farnesene, the aphid alarm pheromone, deterred the flies from laying only at the extremely high dose of 32 mg/plant.
4. Although M. persicae secreted large (1 ng/insect) amounts of alarm pheromone and B. brassicae extremely small (<0.01 ng/insect) amounts, both aphids equally deterred D. radicum from laying.
5. The deterrent effect appeared to result from the aphids physically disturbing the flies during host-plant selection.     

Arabidopsis thaliana leaf-surface extracts are detected by the cabbage root fly (Delia radicum) and stimulate oviposition

Abstract. Oviposition of the cabbage root fly, Delia radicum (Diptera, Anthomyiidae) is stimulated by leaf‐surface extracts of Arabidopsis thaliana (L.) Heynh. (Brassicaceae) ecotype Columbia. The leaf surface of A. thaliana, similar to that of many other crucifers, contains glucosinolates and CIF (‘cabbage identification factor’; 1,2‐dehydro‐3‐thia‐4,10,10b‐triaza‐cyclopenta[.a.]fluorene‐1‐carboxylic acid). These compounds stimulate receptor neurones of the tarsal sensilla of D. radicum whereas additional, unknown compounds are detected by other receptor neurones.     

Orientation behavior of newly hatched larvae of the cabbage maggot,Delia radicum (L.) (Diptera: Anthomyiidae), to volatile plant metabolites

Newly hatched larvae of the cabbage maggot, Delia radicum(L.), responded to various olfactory stimuli from their host plants and oriented themselves by the concentration gradients of these stimuli. Allyl and ethyl isothiocyanate (NCS), characteristic metabolites of host Brassicaceae, elicited positive taxis, but at a higher concentrations were repellent. Benzyl NCS was neutral; only at the highest concentration was it slightly repellent. Phenyl, cyclohexyl, and butyl NCS were repellent in the highest amounts used. Volatile compounds emanating prevailingly from surface parts of plants were both attractive (hexanol, hexanal, cis-3-hexen-1-ol, linalool) and repellent (hexylacetate, cis-3-hexenylacetate, benzaldehyde, myrcene, terpinene, -pinene, limonene). While the attractiveness was associated with NCS group and with unsaturated and saturated alcohols and aldehydes with an optimal sixcarbon chain length, the repellency was probably caused by a cyclized carbon chain and an acetate group. The activity of a compound was strongly influenced by its concentration.     

Ultrastructure of the funicular sensilla of the cabbage root fly,Delia radicum L. (Diptera : Anthomyiidae)

A transmission electron microscope study of the funicular sensilla of the cabbage root fly, Delia radicum, (Diptera : Anthomyiidae), showed 4 types of surface sensilla and 5 types of pit sensilla. The ultrastructure of the surface sensilla indicated all had a primary olfactory function. These include thick-walled multiporous trichoid sensilla, thin-walled multiporous basiconic sensilla (with 2 subtypes), thin-walled multiporous clavate sensilla, and grooved sensilla with channels at the base of each of the grooves. Clavate sensilla had 2 types of dendrites, one tubular, the other “scrolled”. This 2nd type may indicate an additional thermosensitive function. The dorsal pits contained thin-walled multiporous basiconic sensilla with a tapered tip. The ventral pits contain 3 types of sensilla, which have no wall pores and an inflexible socket. These may contain thermo- and/or hygroreceptors and include smooth-walled conical-, smooth-walled tapered- and striated pit sensilla. The 4th type is a grooved pit sensillum similar to the surface type.     

The relationship between cabbage root fly (Delia radicum) larval feeding and the freeze-dried matter and sugar content of Brassica roots

Five genotypes of swede (Brassica napus var. napobrassica), two genotypes of kale (B. oleracea var. acephala), and two genotypes of rape (B. napus var. napus) were each inoculated at the 8–10 true leaf stage with five cabbage root fly (Delia radicum) eggs. The percentage pupation after larval feeding on individual plant genotypes ranged from 45 to 78%, and the mean pupal weight from 6.5 to 13.0 mg. After 5 weeks, larval feeding damage had reduced root weight by up to 47%, compared with uninoculated plants. The dry matter content of undamaged roots was higher in the kales and rapes than in the swedes. Whilst the dry matter content of the rapes and swedes were not changed by D. radicum damage, that of the kales was elevated. The ethanol-soluble sugar content of the root was reduced in all cases by D. radicum larval damage. However, the effect of D. radicum damage on the concentrations of individual sugars (glucose, fructose and sucrose) was crop- and genotype-dependent. In the roots of kales and rapes, the glucose and fructose concentrations were either very low or unaffected by D. radicum damage, whilst both glucose and fructose were generally reduced in swede roots by D. radicum damage. The root sucrose concentration was either reduced or not significantly affected by D. radicum damage in all of the crop types tested. The percentage pupation and the mean pupal weight of D. radicum were inversely correlated to root freeze-dried matter content. D. radicum pupal weight was positively correlated with root fructose, glucose and ethanol-soluble sugar contents.     

Morphology of the antennal sensilla of the cabbage root fly,Delia radicum L. (Diptera : Anthomyiidae)

The antennal sensilla of Delia radicum L. (Diptera : Anthomyiidae) were studied by scanning electron microscopy. On the scape and pedicel, grooved socketed bristles and setiferous plaques were found. There are 4 types of surface sensilla on the funicle: trichoid, basiconic, clavate, and grooved. Their numbers and distribution are described. There are 3–4 single-chambered pits on the dorsal surface of the funicle of both sexes and these contain basiconic sensilla. On the ventral surface, there is one multi-chambered pit, which contains 5 types of sensilla: grooved s., smooth-walled conical s., smooth-walled tapered s., striated s. and a novel type, flattened sensilla. These results are compared with previously published studies on several other fly species.     

Changes in flight track angles of cabbage root fly, Delia radicum, in diffuse clouds and discrete plumes of the host-plant volatile allylisothiocyanate

Flight tracks of female cabbage root fly, Delia radicum (L.), in odourless air, and in difuse clouds and discrete plumes of the host-plant volatile allylisothiocyanate (ANCS), were investigated. In diffuse ANCS D. radicum flew upwind with a smaller mean straight leg track angle with respect to the wind than in odourless air. In discrete ANCS, a larger mean straight leg track angle and a larger range of track angles occurred than in diffuse odour. The frequency distribution of these angles may have been bi-modal. It is suggested that perceived changes in odour concentration may alter the flight track angle. This behaviour is compared to the odour-modulated anemotaxis of male moths locating pheromone sources.

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Résumé Changements des angles de vol de Delia radicum en présence de nuages diffus ou de panaches discontinus d'allylisothiocyanate, substance volatile émise par la plante-hôte.Nous avons étudié les chemins de vol des D. radicum femelles, en air pur, avect des émissions diffuses ou dicontinues de l'allylisothiocyanate (ANCS), substance volatilede la plante-hôte. En présence d'ANCS diffus, D. radicum vole contre le vent. Ce vol présente des portions droites faisant un angle avec la direction du vent; l'angle moyen de ce crochets est plus petit que celui observé en air pur. En présence d'ANCS discontinu, l'angle moyen est plus grand qu'en atmosphère diffuse et surtout l'histogramme de valeur de ces angles est bimodal.On peut envisager que la perception de changements dans la concentration de l'odeur peut modifier l'angle de vol. Un tel comportement est à comparer à l'anémotaxie contre le vent, modulée par l'odeur, observée chez les papillons mâles décelant une source de phéromones.

     

Tarsal chemoreceptors and oviposition behaviour of the cabbage root fly (Delia radicum) sensitive to fractions and new compounds of host-leaf surface extracts

Abstract. Contact chemoreception plays a decisive role in host selection and oviposition behaviour of the cabbage root fly, Delia radicum L. (Diptera, Anthomyiidae). Glucosinolates (mustard oil glucosides) are known to be perceived by the flies, and when sprayed on paper leaf-models induce oviposition. Recently it has become clear that other non-volatile types of compounds must also be involved in host selection.
A pair of ventro-medial C sensilla on die fifth tarsomere respond strongly to a novel compound called tentatively 'cabbage identification factor' (CIF), but not to sucrose, glucose, fructose and proline. CDF is a new non-glucosinolate oviposition stimulant. A single neurone in each sensillum is activated by this compound and the same is true for glucosinolates. In some flies a mixture of bom types of stimuli evoked an apparent mononeural spike train, whereas in odiers spikes of two separate cells were activated. The significance of this variability is not yet clear.
The new stimulant, CIF, does not evoke responses in glucosinolate receptors in the D sensilla. The involvement of the C₃ sensilla in the detection of host-specific compounds constitutes the first known function for C sensilla in D. radicum. CIF appears to be present in leaf surface extracts from the host-plant Brassica oleracea in quantities as low as 1 ng per gram leaf. In spite of this low level, it stimulates oviposition significantly better than glucobrassicin at higher concentrations, which up till now was known as the most powerful stimulant for D.radicum.     

Life history comparison between two competitive Aleochara species in the cabbage root fly, Delia radicum: implications for their use in biological control

This study evaluates the efficacy of Aleochara bilineata Gyll and Aleochara bipustulata L. (Coleoptera: Staphylinidae) as biological control agents against the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae). Biological and demographic characters are documented and comparisons made between the two species. The following results were obtained:(1) The reproductive potential and longevity of A. bipustulata females are significantly higher than those of A. bilineata. These differences are correlated with the biology and ecology of the hosts of both species.(2) Under laboratory conditions the rate of increase (r_m) of both staphylinids species is significantly higher than that of the pest and their major competitor Trybliographa rapae West. (Hymenoptera:Figitidae). This may indicate that A. bilineata and A. bipustulata can be effective biological control agents.(3) Aleochara bilineata appears to be the most interesting biological control agent against D. radicum, because of its higher rate of increase, higher host specificity and host acceptance, and a development time which is well synchronised with that of its host. With its higher competitive ability, a population of A. bilineata may increase even if competition occurs. However, A. bipustulata presents interesting biocontrol characteristics, but the best strategy may be only to use the potential of its natural populations.     

Isolationin vitro ofStrongwellsea castrans [Fungi: Entomophthorales] a pathogen of adult cabbage root flies,Delia radicum [Dipt.: Anthomyiidae]

The entomogenous fungusStrongwellsea castrans was isolatedin vitro for the first time, by incubating conidia projected from infected cabbage root flies (Delia radicum) in a simple, semi-defined liquid medium comprising dextrose, yeast extract and lactalbumin hydrolysate buffered to pH 7. The fungus grew as long unitunicate hyphae. After transfer to a solid nutrient medium, multinucleate hyphal bodies were formed which developed a thick, laminated wall. Neither conidia nor resting spores developed in liquid or on solid media and the fungus survived successive sub-culturing only in liquid media. Using the API-ZYM system, tests on extracts on hyphae ofS. castrans were positive for 11 enzymes but there were no consistent differences in enzyme profiles betweenS. castrans and fungi of the related genusErynia.      

Preference for plants damaged by conspecific larvae in ovipositing cabbage root flies: influence of stimuli from leaf surface and roots

In laboratory dual-choice assays females of the cabbage root fly, Delia radicum, prefer for oviposition plants with roots damaged by conspecific larvae to undamaged controls. Cauliflower and kale plants were inoculated with root fly eggs (25 per plant) and the hatching larvae were allowed to feed on the roots for various periods of time (1–17 days). After 4 (cauliflower) or 5 (kale) days of larval feeding the oviposition preference was most pronounced and flies laid between 64% and 68% of their eggs near plants with damaged roots. Later, with increasing damage but fewer surviving, and thus actively feeding, larvae, the magnitude of the preference declined. The preference for plants already damaged by conspecific larvae may contribute to the previously observed aggregated distribution of D. radicum eggs in Brassica crop fields.Further experiments revealed that the sensory cues inducing this oviposition preference originate from the complex consisting of the damaged roots, the surrounding substrate (soil) and associated microbes, rather than from the aerial plant parts. In choice assays using the root-substrate complex of damaged and control plants (aerial parts removed), the observed preference for damaged roots was similar to that found for the entire plant but was more pronounced. The damaged roots alone, compared to control roots, received up to 72% (cauliflower) and 75% (kale) of the eggs. By contrast, surrogate leaves sprayed with methanolic leaf surface extracts from the most preferred plants which had been damaged were not discriminated from surrogate leaved sprayed with extracts of the respective control plants. Analysis of glucosinolate levels in methanolic leaf surface extracts revealed that root damage resulted in enhanced concentrations of indole-glucosinolates on the leaf surface in kale but not in cauliflower. Although indole-glucosinolates are oviposition stimulants for the cabbage root fly, the induced changes were apparently too small to influence oviposition behaviour.     

The effect of colour of the substrate on the landing and oviposition behaviour of the cabbage root fly

Field and laboratory experiments were made in order to understand the relation between the spectral characteristic of a substrate and its attractiveness for Delia radicum (L.) (Diptera: Anthomyiidae) flies to land or oviposit. Landing females preferred substrates with high reflectance of green and particularly of yellow wavelengths, but substrates with a high proportion of the blue and green reflectance simultaneously (light blue and white) were also very attractive. Unattractive substrates had either low reflectance across the whole insect-visible spectrum (dark blue) or increased reflectance only in orange or red region of the spectrum (red) or a large proportion of UV reflectance (aluminium). Landing males were most attracted to the substrates with the highest total reflection. Oviposition attractiveness of a substrate grew with an increase in the proportion of reflectance in the blue and a decrease in the green regions of the spectrum. In addition, the oviposition attractiveness increased with increasing total reflection and contrast with the background, and decreased with a high proportion of UV reflectance.     

Insect-crop interactions in a diversified cropping system: parasitism by Aleochara bilineata and Trybliographa rapae of the cabbage root fly, Delia radicum, on cabbage in the presence of white clover

Parasitism of the cabbage root fly, Delia radicum (L.) by the staphylinid Aleochara bilineata Gyllenhal and the cynipid Trybliographa rapae Westwood was examined in a cabbage monoculture and a mixed stand of cabbage undersown with white clover. Number of overwintering cabbage root fly pupae per plant was consistently reduced in the mixed stand, and the incidence of plants attacked by cabbage root fly was either reduced or not different in the mixed stand compared to cabbage monoculture. For both parasitoids, the probability of D. radicum attacked plants having at least one parasitized pupa increased with density of cabbage root fly pupae around the plant. For A. bilineata, this positive relation between presence of parasitism and host density was consistently stronger in cabbage monoculture than in cabbage undersown with clover. Location of a host plant by T. rapae was not consistently affected by the presence of clover. D. radicum attacked plants situated in the cabbage and clover mixture were found by T. rapae as easily as in cabbage monoculture. Overall, the total risk of parasitism for a cabbage root fly pupa by A. bilineata was reduced in the mixed stand compared to the cabbage monoculture, whereas the risk of parasitism by T. rapae was not consistently affected by clover. For both parasitoids, intensity of parasitism showed a variable relationship with host density on individual plants attacked by the cabbage root fly. Overall, in spite of consistently lower total density of pupae in the mixed cabbage—clover than in cabbage monoculture, the density of unparasitized pupae was reduced by the presence of non-host plants only in two of the four experiments. The results emphasize the need to include not only herbivore and crop, but also other plant species as well as natural enemies when evaluating management methods.