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.     

The responses of carrot fly larvae, Psila rosae, to components of their physical enrivonment

Abstract. 1. The responses of third instar Psila rosae (F.) larvae to light, temperature, humidity and soil moisture were investigated in the laboratory.
2. Larvae were photonegative and preferred a temperature of about 15°C. Temperatures between 30 and 40°C adversely affected movement and over 40° C were lethal.
3. In choice chambers, larvae preferred humidities of 70–100% r.h. and larvae in sand avoided dry conditions (2.5% field capacity). The latter response became more marked as larvae approached the pre-pupal stage when moistures of 40% field capacity and lower were avoided.
4. Most larvae were found at a depth of 8 cm in sand of uniform moisture content and temperature, but variation in moisture content could alter this preference.
5. In August, most larval damage in the field occurred near the tip of the carrot tap root but was more evenly distributed over the roots in November. It is uncertain whether this was due to soil near the surface being drier in August or whether it was caused by behavioural differences between the two generations of carrot fly larvae.
6. During the summer of 1975, low soil moisture levels resulted in the total absence of larval mines on the carrot roots even though pupae were found at depths of 20–30 cm in the soil. Temperature had no effect on the distribution of mines on carrot roots except at the top 2 cm of the soil profile.     

Oriented responses of carrot fly larvae, Psila rosae, to plant odours, carbon dioxide and carrot root volatiles

ABSTRACT. The responses of carrot fly larvae, Psila rosae , to host plant volatiles were studied. Two criteria for an oriented response were used: one based on whether an individual reached within 1.5 cm of the odour source and one involving analysis of the track patterns of larvae within 3 cm of the odour source. All larval instars were attracted to both host and non-host plant material, though to a lesser degree by the latter; they were also attracted to CO₂, but not to N₂. A steam volatile fraction of carrot roots, and methyl eugenol, a volatile component of this fraction, also attracted the larvae, differentially at different concentrations. The possible roles of both CO₂ and the host-specific secondary plant volatiles in the host plant finding behaviour of soil inhabiting arthropods are discussed.     

Rhythmic activity of adult carrot fly, Psila rosae

ABSTRACT. In a laboratory study of adult carrot fly activity, observations were made on caged populations and on individuals isolated in actographs. In cages in LD 12:12, feeding, oviposition, flight and possibly mating were all rhythmic. Peak feeding occurred within 1 h from lights-on and peak oviposition and flight 10–12 h later. Mating frequency increased slightly in the latter half of the light phase. In actographs in LD 12:12, peak activity also occurred 10–12 h after lights-on. Activity increased with age up to at least 5 days old, independently of the effects of starvation, and by day 5 males were approximately 3 times and females 8 times more active than on the day of emergence. Overall, males were approximately 1.4 times more active than females. Females isolated in actographs for 4 days in continuous dim light showed peaks of activity at intervals of approximately 22.5 h, and activity therefore appears to be a true circadian rhythm. In continuous bright light the activity of females became arrhythmic.     

Relationship between root volatiles of some carrot cultivars and their resistance to the carrot fly, Psila rosae

Field experiments investigated the resistance of some carrot cultivars to Psila rosae. In addition, headspace vapour and steam distillate from the roots of resistant and susceptible varieties were compared by gas-liquid chromatography. The field data confirmed that resistance may operate by decreasing the numbers of eggs laid indicating a nonpreference by the female Psila. Root resistance to the larva was also confirmed but the mechanism was unclear. A new finding was that root resistance is independent of the effect of egg laying, some cultivars evincing one or the other effect and some such as Regulus Imperial displaying both. It was clear that root resistance to the larva is the crucial prerequisite in breeding resistant varieties.One consistent difference was detected by the chemical comparisons: intact roots of resistant varieties released substantially less volatiles. Specifically, Regulus released almost five times less of the volatiles already shown to positively influence host-finding behaviour by the larva.

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Résumé La résistance à P. rosae de quelques cultivars de carotte a été étudiée en plein champ. Parallèlement, les substances volatiles diffusées et celles extraites par la vapeur des racines de variétés résistantes et sensibles, ont été comparées en chromatographie gaz-liquide (GLC). Les résultats en champ ont confirmé que la résistance peut être due à une diminution du nombre d'oeufs pondus, révélant une absence d'attractivité pour les femelles de P. rosae. La résistance des racines aux larves a été aussi confirmée, mais les raisons n'en étaient pas claires. Un aspect nouveau est que la résistance des racines est indépendante de l'effet de la ponte, quelques cultivars présentant l'un ou l'autreeffet et certains, comme Regulus Imperial, manifestant les deux. Il est net que la résistance racinaire aux larves est la condition essentielle pour la sélection de variétés résistantes. Une différence importante a été mise en évidence par les comparaisons chimiques: les racines intactes de variétés résistantes libèrent nettement moins de substances volatiles. Précisément, Regulus a libéré 5 fois moins de substances volatiles déjà connues comme influençant positivement le comportement de découverte de l'hôte par la larve.

     

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.     

Bionomics of the carrot fly Psila rosae F.; the infestation and sampling of carrot crops

The headlands of a carrot field are generally more heavily attacked by carrot fly larvae than the remainder of the field.
The methods of sampling a carrot crop are outlined: it has been found that, for fields up to 10 acres, samples of carrots taken in alternate twos and threes in the midfield, and in twos on the headlands, give a reliable measure of the infestation. For larger fields the number of carrots is doubled. Transects of headlands and whole fields have been made, and are useful in showing the actual amount of damage in a crop. Thus it is possible to represent each field as a three-dimensional body showing the amount and distribution of the attack.
The deterioration of attacked carrot crops during autumn and winter has been followed. Deterioration is measured by (1) percentage carrots attacked, (2) number of mines per 100 carrots and (3) the percentage of carrots unsaleable or unfit for market. The relations between (1) mines per 100 carrots and time, (2) percentage attack and time, (3) mines per 100 carrots and percentage attack, and (4) mines per 100'carrots and percentage unsaleability are stated. Based on the above relationships, methods of prediction of deterioration have been worked out.
The importance of shelter in determining the degree and position of attack is discussed and the relative effects of different types of shelter on infestation are compared.     

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.     

Responses of carrot fly larvae, Psila rosae, to the odorous and contact-chemostimulatory metabolites of host and non-host plants

ABSTRACT. The responses of third instar carrot fly larvae, Psila rosae , to plant metabolites were studied in a moving air olfactometer. Directed responses over distances greater than 7.5 cm were obtained to both host and non-host plant odours, to CO₂ and methyl eugenol, a volatile secondary metabolite of carrot roots. Larvae also oriented to gradients of contact-chemostimuli. The role of both olfactory and contact-chemostimuli in the host plant finding behaviour of P. rosae larvae is discussed, and observational evidence for the behavioural mechanisms involved presented.     

Host-plant susceptibility to the carrot fly, Psila rosae. 1. Acceptability of various host species to ovipositing females

A preference hierarchy of the oligophagous carrot fly for 30 umbelliferous host-plant species or varieties and six non-hosts was assessed. Foliage of the test plants was presented to a laboratory population of the fly in oviposition choice experiments together with leaves of a standard plant, the susceptible carrot cultivar “Danvers”. Only two species (Anthriscus cerefolium, Carum carvi) were more acceptable than the standard plant, while about half the species received significantly fewer eggs. Some umbellifers (Anethum graveolens, Pimpinella major) had a low acceptability similar to that of non-hosts (non-umbelliferous plants). The results obtained with dual and multiple choice assays were in agreement. A multiple choice assay with potted plants yielded a similar ranking of the species as assays using detached leaves, indicating that foliage is representative for whole plants. Variability in the exploratory runs performed by the females on the leaves prior to egg-laying is described and quantified for hosts and non-hosts. Differences in post-alighting pre-ovipositional behaviour of the female flies on the leaves accounted for a major part of the variation recorded in egg-laying.     

Bionomics of the carrot fly (Psila rosae Fab.): II. Soil populations of carrot fly during autumn, winter and spring

The behaviour of the carrot-fly population in the soil during autumn and winter has been investigated in East Anglia over the three seasons 1943-5. The results show that the pest overwinters in both the larval and pupal conditions, and that the proportion of the individuals in each stage varies from crop to crop and from season to season. Samples taken from a May (1943) sown plot showed that pupation, which was 452% complete in October, rose to 742% in March and was not complete until late April. From a June (1944) sown plot of carrots samples showed a rise in percentage pupation from 8-6% in October to 935% in March, the process probably being complete by early April. The changes in proportion of the soil larvae, carrot larvae and puparia in both the May and June plots were followed and are shown graphically. The changes in the size groups of the soil larvae and in the carrot larvae were also traced. The immature larvae persist longer in the carrots than in the soil. The size composition of the larvae from the soil of the June (1944) plot was recorded over the period September to April and the results are given graphically. Samples taken in October 1945 contained highly organized puparia from which flies emerged in a few days. These flies (third generation) gave rise to progeny which overwintered in the larval condition. The population in December 1945, composed chiefly of second generation puparia and third generation larvae, was very similar to that occuring in the previous two years. Mortality in the overwintering carrot-fly larvae and puparia varied considerably but was frequently heavy. The principal agencies concerned were an hymenopterous insect, Dacnusa gracilis Nees., and fungus and bacterial diseases. Other insect parasites occasionally important were Loxotropa tritoma Thorns, and Aleochara sparsa Heer. Nematode parasites were also recorded.     

Problems in field selection for resistance to carrot fly (Psila rosae) in carrot cv. Long Chantenay

In field trials between 1973 and 1982 families of carrots of cv. Long Chantenay were assessed for their resistance to carrot fly (Psila rosae). In some but not all years, families selected for their observed resistance in earlier trials seemed more resistant than the parent line. In more than half the trials there was an association between carrot fly damage and root density, there being less damage on those families from which more roots were harvested. Thus, family selection within cv. Long Chantenay did not lead to a consistent improvement in resistance to carrot fly. Alternative strategies for achieving plant resistance to this pest are discussed.     

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.     

Using physical barriers to prevent carrot fly (Psila rosae (Fabricius)) damage in domestic production

A field experiment was used to assess the effectiveness of different barriers in protecting garden‐scale carrot production from carrot fly (Psila rosae (Fabricius)) damage. Some of the vertical barriers tested were found to provide a useful method of protecting early season carrots from carrot fly in terms of the percentage of carrots free from damage but, under cumulative pest pressure of several generations of carrot fly, such barriers were found to provide insufficient protection. Gardeners should therefore completely cover their carrot crop to attain an acceptable level of control, and this was found to be especially important for carrots harvested later in the season. There were positive effects of some barrier types on yield which may be due, at least in part, to the protection given by the barriers to carrot seedlings.     

Host-plant susceptibility to the carrot fly, Psila rosae. 3. The role of oviposition preferences and larval performance

The acceptability of various plant species to ovipositing carrot flies was weakly, but significantly correlated with the host's suitability for larval development. Both adult host-plant preferences and larval performance as determined in laboratory experiments explained a part of the variation in susceptibility among the various test plants observed in the field. Across the whole set of plant species examined, antixenosis contributed more substantially to resistance than antibiosis, while the reverse seemed to be true for carrot cultivars.     

The influence of plant density and position in field trials designed to evaluate the resistance of carrots to carrot fly (Psila rosae) attack

The results of field trials designed to evaluate the resistance of carrots to carrot fly (Psila rosae) attack were influenced by plant density and position within trials. Five trials are described, their results analysed and implications for future work discussed. Density effects were of major importance when the range of densities within a trial was greater than about three-fold. Density and damage were associated, carrot cultivars and families with the most roots having the least damage; plant densities achieved should therefore be as close as possible to the targets set. Positional effects were often very important, so trials should ideally have no more than about 10 plots in a block, possibly by using an incomplete block design. Conventional analyses of variance removing block effects may be sufficient but it is desirable, especially with large blocks, to use some form of nearest neighbour analysis for which the various possible techniques gave similar results.     

The complementary effects of plant resistance and the choice of sowing and harvest times in reducing carrot fly (Psila rosae) damage to carrots

The benefits of combining a partially-resistant carrot cultivar with different sowing and lifting dates to reduce carrot fly, Psila rosae, damage were investigated at Wellesbourne in 1983 and 1984-85. The partially-resistant cv. Sytan was less damaged and supported fewer insects than the susceptible cv. Danvers on all lifting dates. The estimated reduction of carrot fly larvae on Sytan compared with Danvers ranged from 33 to 95%. Nine combinations of sowing and lifting dates provided more than 75% marketable roots of Sytan compared with only three combinations of dates for Danvers. An early June sowing of both cultivars provided roots of a marketable size with the least attack. More than 90% of Sytan roots were still marketable in December and fewer insects were produced by the end of the season on these roots than on those sown earlier. In addition, sowing in June decreased the number of pupae produced on cv. Danvers by 10 times compared with earlier sowings. Combining partial resistance with specific sowing and lifting times enabled satisfactory yields of marketable carrots to be obtained in a field infested by high populations of carrot fly.     

Electroantennogram responses of the stable fly, Stomoxys calcitrans, to components of host odour

Abstract. Electroantennograms (EAGs) were recorded from laboratory-reared male and female Stomoxys calcitrans (L.) in response to a range of synthetic chemicals known to be electrophysiologically-active for other biting flies. Of the eight compounds initially tested, only two - 1-octen-3-ol and 3-methylphenol - consistently elicited larger electroantennograms (EAGs) than did control treatments; 1-octen-3-ol was the most potent. EAG recovery time was inversely correlated with EAG amplitude. EAGs recorded with primary C₂-C₁₂ carbon chain-length primary aliphatic alcohols peaked at octan-1–ol with pentan-1-ol, hexan-1-ol and heptan-1-ol also eliciting EAG responses significantly larger than the controls. When different C₈ carbon chain compounds and nonane were tested: 1-octen-3-ol elicited the largest EAGs followed by, in decreasing activity, octan-1-ol, 1-bromooctane, octan-3-ol, octanal, 2-octanone, octanoic acid and nonane. The EAG response of 1-octen-3-ol increased sigmoidally with dose, with the threshold at between 2 and 20 ng, and the peak response at 200 μg on filter paper. EAGs larger than control were also elicited by entrained ox odour and ox breath. The behavioural implications are discussed.