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.     

The consistency of the resistance of eight carrot cultivars to carrot fly attack at several centres in Europe

Eight carrot cultivars representing the range of resistance to carrot fly damage as assessed in England were grown at 12 sites in five European countries in 1977 and in 1978. The trials provided evidence of agreement with the findings in England for both years in the ranking of cultivars with respect to their resistance to carrot fly damage. The Nantes cultivar Clause's Sytan Original was the most resistant cultivar tested. Consistent results were obtained from sites where carrots were assessed at harvest and carrot fly attack was severe. A significant interaction between sites and cultivars was largely due to sites where carrots were stored prior to assessment.     

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.     

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.     

Development of carrot inbreds with resistance to carrot fly using a single seed descent programme

Two carrot cultivars, ‘Sytan’ and ‘Long Chantenay’, representing commercially important carrot types and selected for their partial resistance to the carrot fly (Psila rosae) were crossed as the basis for a single seed descent programme. The resulting F₁ progeny were mass pollinated to produce an F₂ generation and approximately 2000 plants were raised from this segregating family in the glasshouse in 1981. By careful choice of sowing date and glasshouse temperatures it was possible to stimulate the plants to flower within 10 months. Individual king umbels were enclosed within bags and pollinated with blowflies. Resulting seed was sown in pots in the following August and the process of seed production repeated in a 12 month cycle. Each inbred line was selfed in this way over three generations until the F₅ stage. Stocks of seed were then multiplied. A total of 753 inbreds were produced by 1987. The vigour of each inbred was evaluated in glasshouse tests and the level of chlorogenic acid was determined by a fluorescence technique. The most vigorous lines with the lowest levels of chlorogenic acid were tested in field experiments against carrot fly at Wellesbourne. Nine inbreds with promising agronomic quality and moderate resistance to carrot fly were selected and seeded. These nine lines were submitted to seed companies with the aim of developing new cultivars of carrot.     

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.     

Preliminary studies of carrot susceptibility to carrot fly attack

Eleven carrot cultivars were tested to compare their susceptibility to carrot fly attack at Wellesbourne and at Mepal, Cambridgeshire in 1969. Significant differences in the percentages of unattacked roots were observed but not all the cultivars behaved consistently in this respect at the two sites. Foliage height and the percentage unattacked roots were not significantly correlated for these cultivars. Four of the cultivars representing the different levels of susceptibility were retested in fen and in mineral soil in microplots at Wellesbourne in 1970. Soil type significantly affected plant growth but not the relative susceptibilities of the cultivars to carrot fly attack. Plant size was positively correlated with insect attack and accounted for most, but not all, of the differences in susceptibility between the cultivars. In 1971 also, cv. Royal Chantenay was significantly less attacked than Speed's Norfolk Giant from August to December even after allowing for the large differences between the two cultivars in plant size. The experiments demonstrated a consistent difference in the relative susceptibilities of cvs Royal Chantenay and Speed's Norfolk Giant. The difference was correlated with plant size but allowances for foliage and root size failed to account for all of the difference. The results suggested a more fundamental basis for part of the difference in the susceptibility of the cultivars to carrot fly attack.     

Further studies relating chlorogenic acid concentration in carrots to carrot fly damage

The effects of first generation carrot fly larval damage on chlorogenic acid concentration in carrots was investigated in a field experiment at Wellesbourne in 1985. In a separate experiment carrots grown in the absence of a resident population of carrot fly were also analysed for chlorogenic acid; these carrots maintained low concentrations of chlorogenic acid through the summer and autumn until low ground temperatures occurred from November to January. The relationship between chlorogenic acid concentration and damage by the first generation of carrot fly was described by a similar model to the one derived previously for late-generation damage but without the cultivar dependence. This may have been because first generation damage takes place in mid-summer when soil temperature is not sufficiently low for differential chlorogenic acid production by carrot cultivars. The model supports the hypothesis that carrot fly damage increases chlorogenic acid production which subsequently encourages further attack. The increase in acid production due to the low winter temperature may be the mechanism which, in turn, induced a differential cultivar response in carrots harvested during the winter.     

The effects of time of sowing and harvest on carrot biochemistry and the resistance of carrots to carrot fly

Two carrot cultivars which represented contrasting levels of resistance to carrot fly, Psila rosae, were grown in a replicated field experiment at Wellesbourne in 1984–85. Seed was sown on five occasions between April and August 1984 and the crop harvested after a minimum growth in the field of 16 wk on seven occasions between August 1984 and February 1985. A highly significant correlation (r = -0.78; P > 0.001) between the levels of carrot fly damage and the concentration of chlorogenic acid in the roots enabled a model to be formulated to predict the relative predisposition to carrot fly larval damage; high levels of chlorogenic acid early in the season predisposed roots to severe attack when they were older.     

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.     

Relationship between the concentration of chlorogenic acid in carrot roots and the incidence of carrot fly larval damage

Chlorogenic acid (1·24-3·36 mg/g) was identified as the main phenolic component in the peel of carrots by hplc analysis. The higher the concentration of chlorogenic acid in different cultivars the greater the susceptibility to carrot fly larval damage. Increases in concentration were found both after carrot fly damage and after carrots had overwintered in the field. The presence and location of chlorogenic acid was confirmed in sections of carrot tissues, mounted in 0·05 M ammonia solution by viewing them using a u.v.-epifluorescent microscope. The importance of phenolic compounds and their function in the production of insect cuticle is discussed in relation to the different concentrations of chlorogenic acid and resistance to carrot fly in carrots.     

An investigation of the host range of the carrot fly

The historical records of the earliest reports of umbelliferous and other hosts of carrot fly ( Psila rosae ) are presented and their validity discussed. In addition, the results are reported of tests made between 1972 and 1980 on 27 species of Umbelliferae and three sub-species of Daucus carota to determine whether they would support the carrot fly. Techniques were developed for identifying plant species as hosts by determining the number of adult flies that emerged from root and soil samples obtained from plants exposed to carrot fly in the field. Experiments in 1979 and 1980 indicating the importance of co-ordinating the life cycles of plant and insect. The results showed that certain annual species of Umbelliferae were unsuitable as hosts in the autumn when they has flowered and senesced. Valid records could only be obtained for these species by exposing the plants to first generation carrot fly attack. Of the 27 species tested, Anthriscus sylvestris, Ferula communis, and Smyrnium olusatrum failed to support carrot fly and the following 13 species were new host records: Aethusa cynapium, Anthriscus caucalis, Apium inundatum, Apium nodiflorum, Bupleurum tenuissimum, Chaerophyllum temulentum, Cicuta virosa, Daucus capillifolius, Daucus glochidiatus, Heracleum mantegazzianum, Ligusticum scoticum, Oenanthe crocata and Sison amomum . From two species, Chaerophyllum temulentum and Torilis japonica , more than two carrot flies/root were recovered. These species are numerous and widespread in carrot-growing districts. The significance of these and other wild host plants in relation to the ecology and control of carrot fly is discussed.     

Short feeding period of carrot psyllid (Trioza apicalis) females at early growth stages of carrot reduces yield and causes leaf discolouration

Overwintered adult carrot psyllids [Trioza apicalis Förster (Homoptera: Psylloidea: Triozidae)] damage carrot [(Daucus carota ssp. sativum L.) (Apiaceae)] seedlings by phloem feeding on the leaves. The aim of this study was to investigate the carrot root and shoot growth in relation to carrot psyllid density during early growth stages. One, two, or three carrot psyllids were allowed to feed on carrot seedlings for 3 days. Leaf damage was measured at the 8‐leaf stage, and root, leaf fresh weight, and number of true leaves were measured at harvest. Both the age of the carrot seedling at infestation and the psyllid density had a significant effect on leaf damage at the 8‐leaf stage: seedlings damaged at the cotyledon stage exhibited more leaf damage than seedlings damaged at the 1‐leaf stage. A higher psyllid density significantly reduced the carrot root weight at harvest. The significant interaction of psyllid density with seedling age indicates that differently aged carrot seedlings responded differently to feeding: one psyllid feeding for 3 days at the cotyledon stage caused a significant yield loss, whereas three psyllids were needed to cause the same impact at the 1‐leaf stage. Carrot leaf weight at harvest was not reduced by carrot psyllid feeding: leaves recovered from the damage but roots did not. Our results confirm the farmers’ observations that a trap replacement period of 1 week for carrot psyllid monitoring is too long, especially at the cotyledon stage. Severe leaf discolouration on damaged carrots was observed at harvest. The possible reasons for this discolouration, such as toxin excreted in psyllid saliva or plant pathogenic mycoplasma infection, are discussed.     

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.     

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.     

A relationship between dry matter and damage by larvae of the turnip root flyDelia floralis in cultivars of the swedish turnipBrassica napus

A series of trials from 1981–85 investigating the amount of damage caused by larvae of the turnip root flyDelia floralis (Fallen) to cultivars of Swedish turnipBrassica Napus (L.) of varying percentages of dry matter indicated that those of high dry matter were damaged less than low dry matter cultivars. Pupal numbers of turnip root fly gave good correlations with damage but there was no relationship between damage and cabbage root fly pupae.     

A rapid non-destructive technique for screening carrot seedlings for resistance to carrot fly damage

A technique based on fluorescence of carrot root tissues has been devised to enable large numbers of accessions to be screened for resistance to carrot fly using an epi-fluorescence microscope. The technique is non-destructive, and has been partly automated to assess at least 400 seedlings/hour. A data-capture and handling package provides prompt summarising and print-out of the results.     

Effect on carrot root yield and size distribution of the growth-retardants chlormequat chloride and aminozide

In two experiments, foliar sprays of either chlormequat chloride or aminozide depressed top growth, increased root yield and enhanced leaf and root colour in six carrot cultivars. In the first experiment, a direct correlation between concentration of the two retardants and root yield was apparent within the range 0–2000 mg/1 a.i. applied as run-off sprays to two cultivars. In the second experiment, increases in total root and canning-size yield and root/shoot ratio were obtained with four cultivars, but the magnitude of these effects was dependent on the time at which the retardants were applied and the time of harvest of the crop. The effects of aminozide on root yield were less persistent than those of chlormequat chloride, the yield differences between untreated and aminozide-treated plots decreasing with time after application.     

Intensity of radicle fluorescence as related to the resistance of seedlings of lettuce to the lettuce root aphid and carrot to the carrot fly

A technique is described which indicates by u.v. fluorescence the concentration of caffeoylquinic, chlorogenic and isochlorogenic acids in roots of germinating lettuce and carrot. The surface fluorescence of radicles from lettuce root aphid (Pemphigus bursarius) resistant seedlings was more intense than for susceptible seedlings, attributable to the higher concentration of isochlorogenic acid. In contrast, radicles of carrot seedlings resistant to carrot fly (Psila rosae) were less fluorescent than susceptible seedlings, corresponding to the lower concentration of chlorogenic acid in resistant seedlings. The technique is non-destructive and has been developed to distinguish between cultivars or breeding lines potentially resistant to these insect pests.     

A method of using the proportions of undamaged carrots or parsnips to estimate the relative population densities of carrot fly (Psila rosae) larvae, and its practical applications

The plant-to-plant distribution of carrot fly damage was studied on carrots and parsnips in field and microplot experiments. The observed data were well-fitted by the relationship In (–In q) = In L+ constant, where q is the proportion of undamaged roots and L is the number of larval mines in a sample and is proportional to the number of associated larvae, N. Hence, if N₁ and N₂ are the numbers of larvae at two levels of infestation on one occasion, the relative numbers present, N₂:N₁ are estimated by In q₂: In q₁ In this way the relative efficiencies of insecticide treatments can be determined within an experiment from only q₁ and q₂ and compared between occasions. A nomogram relating q₁, q₂ and treatment efficiency is provided. The method can also probably be used to compare the relative susceptibilities of host-plant cultivars to carrot fly attack. Guidance is given on the sample sizes needed to estimate q within 95% confidence limits and constraints on the application of the method are stated. An appropriate statistical procedure for analysing data sets on carrot fly infestations is by analysis of deviance and an example is given in an Appendix. The principles described can probably be adapted to other crop/pest problems.