Behavioural responses of the carrot fly larva, Psila rosae, to carrot root volatiles

ABSTRACT. The behaviour of the carrot fly larva, Psila rosae F. (Diptera, Psilidae), was investigated in the presence of six concentrations of thirty-four volatiles associated with carrot root and of trans -methyl-iso-eugenol, a constituent of carrot herb oil. Bomyl acetate, 2,4-dimethyl styrene, α-ionone, SbT-ionone and biphenyl were the most consistently preferred compounds. Klinotaxis and klinokinesis were the orientation responses near the stimulus source. Trans -2-nonenal was the most consistently avoided compound. Extracts derived by steam distillation of carrot roots and by concentration of vapour from roots were resolved, by gas liquid chromatography, into forty-five and twenty-two major constituents, respectively. The five preferred compounds are micro-components of these extracts, collectively representing 1% and 0.6% of root extract oil and vapour, respectively. It is suggested that these secondary plant constituents are specific host finding cues for the larva, and that frara-2-nonenal is a carrot defence compound.     

Carrot fly cultivar preferences: some influencing factors

Abstract. 1. Factors affecting cultivar preferences by the carrot fly Psila rosae F. were examined in choice tests using foliage from carrot lines representing extremes of resistance and susceptibility to attack.
2. Flies were offered a choice to oviposit on foliage, artificial substrates impregnated with leaf surface extracts or enclosed in vapours of different cultivars to establish the importance of contact and olfactory stimuli. Susceptible cv. Danvers Half Long 126 foliage was preferred over that of resistant cvs. Clause's Sytan Original and Tip-Top. But, whereas the Sytan leaf surface extract was as effective as that of Danvers, the surface extract of Tip-Top was very much inferior as an oviposition stimulant. By contrast, Sytan and Tip-Top foliar vapours were both more attractive to the fly than that from Danvers.
3. Headspace vapours over Sytan foliage evoked significantly higher electroantennogram responses than those from Danvers.
4. Cold-trapped foliar volatiles of Sytan and Danvers were analysed by gas chromatography linked with antennographic detection. Foliage of Sytan released, among other chemo-stimulants, higher levels of host plant attractants, i.e. green leaf aldehydes and the phenylpropanoids, trans-asarone and trans-methylisoeugenol.
5. These findings indicate that olfactory and contact chemostimuli may be involved in preferences shown by the carrot fly for carrot cultivars, but these factors alone do not fully account for the preferences observed.     

Further investigations of the host range of the carrot fly, Psila rosae (F.)

Techniques were developed for testing umbelliferous species against naturally-occurring populations of carrot fly in the field. The most efficient technique involved exposure of plants to either first or second generations of the insect in the field, followed by caging of infested plots and trapping of emerged flies in water in yellow water dishes. In a series of field experiments between 1981 and 1989 inclusive, a total of 132 umbelliferous species and sub-species were tested against carrot fly. Seventy-eight of these proved to be new hosts, 27 were confirmed as hosts and 27 failed to support any carrot flies. Six non-umbelliferous plant species failed to be colonised by carrot fly. Sources of variability in investigations of the host range of insects are discussed.     

Monitoring populations of the carrot fly, Psila rosae, with sticky and water traps

The relative, effectiveness of Rebell®, small cylinder, large cylinder, windmill and water traps, the five types of trap used currently for monitoring populations of the carrot fly, Psila rosae, was assessed in nine field experiments, three in south west Lancashire, four in the Fens (Suffolk, Norfolk, Cambridgeshire), one in East Suffolk and one at Wellesbourne, Warwickshire. Regression analysis of the numbers of flies caught on each type of trap against the numbers caught on the Rebell® trap indicated that each trap samples a constant proportion of the fly population relative to the other traps. Therefore, provided the fly population was sufficiently large for insects to be caught on the least effective traps, any of the five traps would monitor adequately fluctuations in carrot fly populations. However, the Rebell® trap caught 4–17 times as many flies/trap and 5–7 times as many flieshnit area of trap as any of the other traps tested.
Operators considered the Rebell® trap to be the easiest to use. However, it was more expensive than any of the other traps tested.     

Exploitation of the resistance to carrot fly in the wild carrot species Daucus capillifolius

Under field conditions a wild Daucus species from Libya, D. capillifolius, supported less than one tenth as many carrot flies (Psila rosae) as the susceptible carrot cultivar Danvers Half Long 126. Breeding lines developed from crosses between D. capillifolius and three different carrot types were grown in a series of field experiments at Wellesbourne between 1980 and 1989. Each year selections were made for agronomic quality and/or for increased resistance to carrot fly. The programme produced lines which for size, shape and colour represented most of the commercially-important carrot types. Some of these lines were also significantly more resistant to carrot fly than selections from the partially-resistant cv. Sytan. However, the best lines were not as resistant as the wild parent. The highest quality resistant lines were sold to seed companies for variety production.     

Abnormal egg-laying behaviour of female carrot flies (Psila rosae) induced by the fungus Entomophthora muscae

Female carrot flies (Psila rosae F.) infected with Entomophthora muscae (C.) Fres. on days 1 to 4 of adult life were unable to lay their eggs near the food plant. Flies infected on day 5 of adult life showed an egg-laying behaviour similar to that of uninfected flies. E. muscae-infected female carrot flies caught in the field showed an egg-laying behaviour comparable to that of the flies infected in the laboratory during the first day of their live.Eggs laid by infected females were equally as fertile as eggs laid by uninfected females, and developed into normal flies.It is concluded that female carrot flies infected with E. muscae in the field do not in general contribute to the development of the carrot fly population, due to their abnormal egg-laying behaviour.

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Zusammenfassung Wenn Weibchen der echten Möhrenfliegen (Psila rosae) zwischen den ersten und der vierten Tag in dem Laboratorium mit E. muscae infiziert worden waren, könnten sie ihre Eier an den Futterpflanze nicht ablegen. E. muscae-infizierte Weibchen, die ins Feld gefangen worden waren, zeigten auch ein abweichendes Eiablageverhalten. Ihr Behalten war mit dem Verhalten von Fliegen vergleichbar, die an ihrem ersten Lebenstag im Laboratorium infiziert worden waren.Von infizierten Weibchen gelegte Eier waren ebenso fruchtbar wie die von nicht infizierten Weibchen gelegten Eiern. Es wird konkludiert, dass E. muscae-infizierten Weibchen der echten Möhrenfliegen der Entwicklung der Population keinen Beitrag leisten.

     

The culture ofEntomophthora muscae (C) Fres. in carrot flies (Psila rosae F.) and the effect of temperature on the pathology of the fungus

A method for maintaining anin vivo culture ofEntomophthora muscae (C) Fres. on its original host, adult carrot flies (Psila rosae F.), is described. The lethal time for adult carrot flies was greatly influenced by temperature, both for infected and for uninfected flies. In the range 8.2°C–20.2°C the LT₅₀ for infected flies was about 5.4 times shorter than the estimated average life-span for uninfected flies. The discharge of primary spores was also strongly dependent on temperature. The total number of primary spores discharged per fly at 100% RH and in darkness ranged between 1.2×10⁴ and 9.6×10⁴ with a mean of 5.1×10⁴.      

An analysis of the growth response of carrot seedlings to deficiency in some mineral nutrients

Critical plant concentrations for a reduction in relative growth rate to 90% of that of fully nourished plants were estimated by a novel method for several mineral nutrients. Carrot plants were grown from seed for 28 days in a range of nutrient solutions omitting N, P, K, Ca, S, Mg, Fe, B, Mn, Zn, Cu and Mo as separate treatments. All treatments except -Mn, -Zn, -Cu and -Mo resulted in effects on plant growth and the development of deficiency symptoms. Estimates of critical concentrations were based on a simple simulation model incorporating the principle of nutrient dilution with increasing plant weight and on mineral analysis of the plants. Parameters governing the shape of the relationship between fractional relative growth rate and plant nutrient concentration were altered until the model predicted the observed final mean dry weight of deficient plants and time of divergence of this growth curve from that of fully nourished plants. Critical concentrations so obtained were higher than those previously reported for Ca, Fe, N and P in carrots and lower for K, Mg and S.     

Effects of root restriction and growth regulator treatments on the growth of carrot (Daucus carota L.) seedlings

The growth of carrot (Daucus carota L.) seedlings in clear polystyrene vials (7.5 × 2.5 cm diameter) which were either perforated or unperforated and suspended in a large container containing nutrient solution was compared with that of similar seedlings grown directly in the main container. After 28 days plants grown in the perforated, aerated vials had smaller shoots, tap roots (c 20 percent reduction) and fibrous roots (c 60 percent reduction) than unrestricted plants. In the absence of adequate aeration, the effects of root restriction were greater. Foliar applications of ethephon to the leaves of unrestricted plants reduced shoot weights but had little effect on either tap root or fibrous root weights, or even increased tap root weight at lower concentrations. Silver thiosulphate, aminovinylglycine and a highly mobile cytokinin (SD8339) all slightly enhanced shoot growth of restricted plants but had no effect on either tap roots or fibrous root development. It is suggested that although root restriction effects may be partly mediated by endogenous hormones, it seems unlikely that such effects are due solely to stimulation of ethylene production or a reduction in endogenous cytokinin levels.Abbreviations AVG Aminovinylglycine - BA N⁶-Benzyladenine - Ethephon 2-Chloroethylphosphonic Acid - SD8339 6-Benzylamino-9-(Tetrahydropyran-2-yl)-Adenine - STS Silver Thiosulphate     

Host-plant susceptibility to the carrot fly, Psila rosae. 2. Suitability of various host species for larval development

Several apiaceous and two asteraceous species were tested for their suitability to support larval development of the carrot fly. Plants grown in pots or transplanted from seed beds into pots, were inoculated with a specific number of eggs. Pupae and non-pupated larvae were collected 6–7 wk after inoculation. Both the number and weights of pupae produced varied widely among the species. Cultivated carrots Daucus carota sativus often gave rise to only moderate numbers of pupae, but these invariably attained the highest weights. Pimpinella major was the only apiaceous plant tested that did not yield any carrot flies. The two asteraceous plants Cichorium intybus and Tanacetum vulgare failed to support larval development. Total carrot fly biomass produced per plant was influenced by both the host species and the root weight. Emergence rates of adult flies were positively correlated with pupal weights. Small individuals tended to have a longer total developmental time from egg to adult fly.     

Effects of plant odor on oviposition by the black swallowtail butterfly,Papilio polyxenes (Lepidoptera: Papilionidae)

Black swallowtail females laid more eggs on plant models treated with contact stimulants and volatiles from carrot leaves than on models treated only with contact stimulants. The volatiles enhanced landing rates and females alighted more frequently on artificial leaves treated with host volatiles than on adjacent control leaves. Volatiles from cabbage, a nonhost, inhibited landing rates on artificial leaves treated with carrot contact stimulants. Examination of antennae revealed two major types of sensilla, believed to be olfactory in function. Electroantennogram preparations responded more strongly to carrot volatiles than to cabbage volatiles and several shared responses at particular retention times to carrot volatile components eluting from a gas chromatograph. Our results are consistent with a long-standing hypothesis that behavioral responses to essential oil components characteristic of the larval food plants have facilitated host shifts in the genus Papilio.     

The stability of accumulated anthocyanin in suspension cultures of the parental line and high and low accumulating subclones of wild carrot

Abhibition of the accumulation of anthocyanin but not of growth in wild carrot cultures was achieved using 2-amino-oxyphenylpropionic acid or phenylpropiolic acid. The concentration of inhibitor required to achieve this differential effect increased with increased culture density.By several additions of inhibitor at differential times, anthocyanin accumulation in the parental culture and in high-and low-acculmulating subclones could be completely inhibited. Under these conditions the level of anthocyanin in the cultures remained unchanged. These data show that the accumulated anthocyanin is not metabolized and that differences in the rate of degradation of anthocyanin does not account for the differences in anthocyanin accumulation by the subclones examined.     

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.     

Mechanisms of resistance to onion fly egg-laying

When gravid onion fly females, Delia antiqua (Meigen) (Diptera: Anthomyiidae) were presented in laboratory choice test with 6-week-old plants of a susceptible cultivar and of onion breeding lines selected for resistance, mean numbers of eggs laid ranged from 34.8 to 1.6 eggs per plant. Differences in ovipositional responses were mirrored by differences in plant size. Analysis of covariance revealed no significant differences in ovipositional responses to breeding lines when differences in size were taken into account. Foliar surrogates were developed so that single size parameters could be varied while holding all other plant stimuli constant. Tests using these surrogates revealed that among plants with basal diameters of 1 to 4 mm and heights of 100 to 350 mm, diameter alone significantly influenced responses of ovipositing females. Ovipositional responses to plants beyond this size range could not be explained strictly by diameter differences. These results underscore the necessity of considering plant size as well as plant chemistry and texture in efforts to evalute and design resistant cultivars.

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Résumé A la lumière d'opinions récentes concernant la découverte des plantes-hôtes par D. antiqua, nous avons examiné les conditions de la résistance chez plusieurs lignées de A. cepa les moins attaquées lors d'essais en plein champ aux Pays-Bas, et dont on considérait que la résistance était due à une antixénose.Des femelles gravides en présence, au laboratoire, d'un choix de plants de 6 semaines, pondent en moyenne 1,6 à 34,8 oeufs par plant. Les différences entre les pontes correspondent aux tailles des plants. Si l'on tient compte de ces dernières, l'analyse de covariance ne montre pas de différence significative entre les pontes sur les différentes lignées. Des substituts de feuilles ont été créés de façon à faire varier les paramètres de taille tout en maintenant constants les autres stimuli de la plante: pour des plantes, dont le diamètre à la base variait de 1 à 4 mm et la hauteur de 100 à 350 mm, seul le diamètre a significativement influé sur la ponte. Les pontes sur les plantes hors de ces dimensions n'ont pu être expliquées par les seules différences de diamètre.Comme les paysans des principales régions productrices d'oignons d'Europe et des U.S.A. font des bénéfices importants avec la culture d'oignons précoces, ces lignées, sélectionnées par mégarde pour leur développement lent, ne sont probablement pas une bonne source de résistance à la mouche de l'oignon. Cependant, ces résultats soulignent la nécessité de tenir autant compte de la taille de la plante que de son chimisme ou de sa texture lors de l'évaluation et de la création de cultivars résistants.

     

Physical and chemical factors influencing landing and oviposition by the cabbage root fly on host-plant models

Various plant models were used in both choice and no-choice tests in the laboratory to assess landing and oviposition preferences of the cabbage root fly,Delia radicum (L.). The main factor governing the site most suitable for landing was the conspicuousness of the object and not its shape. Oviposition was influenced considerably by the pre-conditioning of the females. Deprived females laid eggs even when denied access to both host plant chemicals and host-plant models. The dominant role of contact chemical stimuli in host acceptance was reconfirmed, but only a combination of physical and chemical stimuli appeared capable of eliciting normal oviposition. The combination of contact chemical stimuli and the presence of a stem on the test model had a synergistic effect on the numbers of eggs laid in both choice and no-choice situations. In choice bioassays, female cabbage root flies distinguished between models of different shapes, heights and sizes. The size and shape of the models appeared to be perceived in part after the flies had landed.     

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.

     

Combined expression of chitinase and lipid transfer protein genes in transgenic carrot plants enhances resistance to foliar fungal pathogens

Two pathogenesis-related (PR) protein genes consisting of a barley chitinase (chi-2) and a wheat lipid-transfer-protein (ltp) were introduced singly and in combination into carrot plants via Agrobacterium-mediated transformation using the phosphinothricin acetyl transferase (bar) gene as a selectable marker. Over 75% of regenerated plants were confirmed to be positive for the transgenes by PCR and RT-PCR and were resistant to the herbicide Liberty (0.2%, v/v). Northern analysis and immunoblotting confirmed the expression of the transgenes in about 70% of the plants, with variable expression levels among individual lines. Southern analysis revealed from one to three copies of each transgene. Transgenic plants were inoculated with two necrotrophic foliar fungal pathogens, Alternaria radicicola and Botrytis cinerea, and showed significantly higher resistance when both PR genes were expressed compared to single-gene transformants. The level of disease reduction in plants expressing both genes was 95% for Botrytis and 90% for Alternaria infection compared to 40–50% for single-gene transformants. The chi2 and ltp genes could be deployed in combination in other crop plants to significantly enhance resistance to necrotrophic fungal pathogens.     

Electroantennogram responses of the oriental fruit fly, Dacus dorsalis, to a spectrum of alcohol and aldehyde plant volatiles

Electroantennograms (EAGs) were recorded from unmated, laboratory-reared, male and female oriental fruit flies, Dacus dorsalis, in response to a range of between C₁ and C₁₂ carbon chain-length saturated and unaturated aliphatic alcohols and aldehydes, most all of which are known host-plant volatiles. Only two of the 35 compounds tested elicited significantly larger EAGs from female than male antennae. For the two functional-group series tested, aldehydes elicited responses greater than or equal to the responses to the alcohols. In general, the unsaturated alcohols did not elicit responses significantly different from the saturated alcohols. However, the unsaturated aldehydes, (E)-2-hexenal and 10-undecenal, elicited larger amplitude EAGs than their saturated analogs. EAGs were significantly greater for a particular carbon chain-length, with responsiveness to primary alcohols peaking at C₆ and aldehydes peaking at C₇. The (E)-2- monoenic alcohols peaked at C₆, while the (E)-3-alcohols plateaued between C₅ and C₈. The greatest EAG responses of all compounds tested were elicited by the saturated and unsaturated C₆ alcohols and aldehydes which are constitutents of the general green-leaf volatile complex that emanates from most plants. The potential adapative benefit of selective sensitivity to green-leaf volatiles is discussed in regards to foraging behaviors of oriental fruit flies.

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Résumé Des électroantennogrammes (EAG) ont enregistré les réponses, en élevages de femelles et mâles vierges de Dacus dorsalis, à une gamme de chaînes de carbones de C₁ à C₁₂ saturés et non-saturés d'alcools aliphatiques et d'aldéhydes, dont beaucoup sont connus comme substances volatiles des végétaux. Seulement 2 des 35 composés examinés ont provoqué des EAG significativement plus importants chez les femelles que chez les mâles. Pour les séries des deux groupes fonctionnels examinés, les aldéhydes ont provoqué des réponses supérieures ou égales aux alcools. En général, les réponses aux alcools nonsaturés n'étaient pas significativement différentes des réponses aux alcools saturés. Cependant, les aldéhydes non-saturés, (E)-2-hexénal et 10-undécénal, ont induit des EAG de plus grande ampleur que leurs analogues saturés. Les EAG étaient significativement les plus importants pour une chaîne de longueur particulière, la réponse aux alcools primaires culminant en C₆ et les aldéhydes en C₇. Les alcools monoéniques (E)-2- culminaient en C₆, tandis que les alcools (E)-3- étaient étales entre C₅ et C₈. Les EAG les plus importants ont été obtenus pour tous les composés examinés avec les alcools et aldéhydes en C₆ qui appartiennent à l'odeur verte complexe émise par beaucoup de plantes. Le bénéfice adaptatif potentiel de la sensibilité sélective à l'odeur verte des feuilles est examinée en fonction du comportement de prospection de D. dorsalis.