Tom W. Pope

Varying responses of insect herbivores to altered plant chemistry under organic and conventional treatments

The hypothesis that plants supplied with organic fertilizers are better defended against insect herbivores than those supplied with synthetic fertilizers was tested over two field seasons. Organic and synthetic fertilizer treatments at two nitrogen concentrations were supplied to Brassica plants, and their effects on the abundance of herbivore species and plant chemistry were assessed. The organic treatments also differed in fertilizer type: a green manure was used for the low-nitrogen treatment, while the high-nitrogen treatment contained green and animal manures. Two aphid species showed different responses to fertilizers: the Brassica specialist Brevicoryne brassicae was more abundant on organically fertilized plants, while the generalist Myzus persicae had higher populations on synthetically fertilized plants. The diamondback moth Plutella xylostella (a crucifer specialist) was more abundant on synthetically fertilized plants and preferred to oviposit on these plants. Glucosinolate concentrations were up to three times greater on plants grown in the organic treatments, while foliar nitrogen was maximized on plants under the higher of the synthetic fertilizer treatments. The varying response of herbivore species to these strong differences in plant chemistry demonstrates that hypotheses on defence in organically grown crops have over-simplified the response of phytophagous insects.

The effect of ant attendance on the success of rosy apple aphid populations, natural enemy abundance and apple damage in orchards

1 The rosy apple aphid, Dysaphis plantaginea, is the most serious pest of apple in Europe and, although conventionally controlled by insecticides, alternative management measures are being sought. Colonies of D. plantaginea are commonly attended by ants, yet the effects of this relationship have received little attention.

Interactions Between Host-plant Volatiles and the Sex Pheromones of the Bird Cherry-oat Aphid, Rhopalosiphum padi and the Damson-hop Aphid, Phorodon humuli

The bird cherry-oat aphid, Rhopalosiphum padi (L.), and the damson-hop aphid, Phorodon humuli (Schrank), migrate at the same time of year and colonize closely related Prunus spp. as primary hosts, but utilize (1R,4aS,7S,7aR)-nepetalactol and (1RS,4aR,7S,7aS)-nepetalactol, respectively, as sex pheromones. Interactions between these sex pheromones and benzaldehyde and methyl salicylate, plant volatiles common to primary hosts of both species, were investigated to assess whether they confer reproductive isolation between these species. Female autumn migrants (gynoparae) and males of these two species were caught in the field with water traps baited with their respective sex pheromones. Rhopalosiphum padi gynoparae and males also responded positively to benzaldehyde. Release of either benzaldehyde or methyl salicylate with the conspecific sex pheromone increased catches of both species of aphid. However, releasing both plant volatiles with the sex pheromone of R. padi increased catches of gynoparae and males, but reduced those with the sex pheromone of P. humuli. These results support the hypothesis that specific plant volatiles synergize responses of autumn migrating aphids to their sex pheromone. Because these interactions are species-specific, they may be important in allowing males to discriminate between conspecific sexual females (oviparae) and those of other aphid species.

Interactions among phytophagous mites, and introduced and naturally occurring predatory mites, on strawberry in the UK.

In choice test experiments on strawberry leaf disc arenas the phytoseiid mites Neoseiulus californicus and N. cucumeris were more effective than Typhlodromus pyri as predators of the phytophagous mites Tetranychus urticae and Phytonemus pallidus. There were no preferences shown for either prey by any of these predators. In multiple predator leaf disc experiments both Phytoseiulus persimilis and N. cucumeris significantly reduced numbers of T. urticae eggs and active stages; this effect was seen when the two species were present alone or in combination with other predator species. Neoseiulus californicus was less effective at reducing T. urticae numbers, and T. pyri was not effective; no interaction between predator species was detected in these experiments. When T. urticae alone was present as prey on potted plants, P. persimilis and N. californicus were the only phytoseiids to significantly reduce T. urticae numbers. These two predator species provided effective control of T. urticae when P. pallidus was also present; however, none of the predators reduced numbers of P. pallidus. There were no significant negative interactions when different species of predators were present together on these potted plants. In field experiments, releases of both P. persimilis and N. cucumeris significantly reduced T. urticae numbers. However, there was a significant interaction between these predator species, leading to poorer control of T. urticae when both species were released together. These results show the importance of conducting predator/prey feeding tests at different spatial scales.

Responses of the aphids Phorodon humuli and Rhopalosiphum padi to sex pheromone stereochemistry in the field.

Gynoparous female and male damson-hop aphids, Phorodon humuli (Schrank), were caught in the field by water traps that were releasing the sex pheromone of this species, (1RS,4aR,7S,7aS)-nepetalactol. No behavioral activity was elicited by (4aS,7S,7aR)-nepetalactone, the major sex pheromone component of other aphid species such as Megoura viciae Buckton, even though olfactory cells were found in the secondary rhinaria on the third antennal segment of P. humuli that responded strongly to this compound. Gynoparous female P. humuli in the field responded less strongly to (1R,4aS,7S,7aR)-nepetalactol, the sex pheromone of the bird cherry-oat aphid, Rhopalosiphum padi (L.), than they did to the (4aR,7S,7aS)-nepetalactols, but males responded only to the latter. The (4aR,7S,7aS)-nepetalactone showed no electrophysiological activity so was not used in field trials. Releasing either the (4aS,7S,7aR)-nepetalactone or the (1R,4aS,7S,7aR)-nepetalactol with the (4aR,7S,7aS)-nepetalactols did not inhibit the response of P. humuli gynoparous females and males to the latter. Males of R. padi responded as strongly to the (4aR,7S,7aS)-nepetalactols as they did to (1R,4aS,7S,7aR)-nepetalactol. Males of P. humuli and R. padi responded positively to an increased concentration of the (4aR,7S,7aS)-nepetalactols released from two vials compared with that from a single vial, as did P. humuli (in one of two experiments) and R. padi to the (1RS,4aR,7S,7aS)- and (1R,4aS,7S,7aR)-nepetalactols when released together.

Effects of organic and conventional fertilizer treatments on host selection by the aphid parasitoid Diaeretiella rapae

The type and quantity of fertilizer supplied to a crop will differ between organic and conventional farming practices. Altering the type of fertilizer a plant is provided with can influence a plant’s foliar nitrogen levels, as well as the composition and concentration of defence compounds, such as glucosinolates. Many natural enemies of insect herbivores can respond to headspace volatiles emitted by the herbivores’ host plant in response to herbivory. We propose that manipulating fertilizer type may also influence the headspace volatile profiles of plants, and as a result, the tritrophic interactions that occur between plants, their insect pests and those pests’ natural enemies. Here, we investigate a tritrophic system consisting of cabbage plants, Brassica oleracea, a parasitoid, Diaeretiella rapae, and one of its hosts, the specialist cabbage aphid Brevicoryne brassicae. Brassica oleracea plants were provided with either no additional fertilization or one of three types of fertilizer: Nitram (ammonium nitrate), John Innes base or organic chicken manure. We investigated whether these changes would alter the rate of parasitism of aphids on those plants and whether any differences in parasitism could be explained by differences in attractivity of the plants to D. rapae or attack rate of aphids by D. rapae. In free‐choice experiments, there were significant differences in the percentage of B. brassicae parasitized by D. rapae between B. oleracea plants grown in different fertilizer treatments. In a series of dual‐choice Y‐tube olfactometry experiments, D. rapae females discriminated between B. brassicae‐infested and undamaged plants, but parasitoids did not discriminate between similarly infested plants grown in different fertilizer treatments. Correspondingly, in attack rate experiments, there were no differences in the rate that D. rapae attacked B. brassicae on B. oleracea plants grown in different fertilizer treatments. These findings are of direct relevance to sustainable and conventional farming practices.

Treating hop plants with (Z)-jasmone increases colonization by Phorodon humuli (Hemiptera: Aphididae) spring migrants

Hop plants were sprayed with (Z)-jasmone, at a rate of 50 g ha(-1), during the spring migration of the damson-hop aphid Phorodon humuli (Schrank) in 2002 and 2003. Numbers of P. humuli spring migrants colonizing hop plants, Humulus lupulus L., 2-6 and 7-11 days after applying this treatment were assessed in both years. During the first five-day period, significantly more spring migrants were found on hop plants treated with (Z)-jasmone, in comparison with control plants, in 2002. By contrast, no significant difference was evident in the second five-day period. Although the migration in 2003 was much lighter than in 2002, greater numbers of migrants were again removed from treated plants. Indeed, more spring migrants were removed from plants sprayed with (Z)-jasmone in this year during both five-day periods (11 and 44%, respectively) compared with the 23% greater numbers removed in the first five-day period in 2002. Therefore, unlike some other species of aphid, where numbers were consistently lower on plots sprayed with (Z)-jasmone, spraying the secondary host of P. humuli with this compound appears to increase colonization by spring migrants.

Development and optimisation of a sex pheromone lure for monitoring populations of saddle gall midge, Haplodiplosis marginata

Saddle gall midge, Haplodiplosis marginata (von Roser) (Diptera: Cecidomyiidae), is a sporadic pest of cereals in Northern and Central Europe and is of increasing importance in the UK. Recently, the major component of the sex pheromone produced by adult female H. marginata was reported to be 2‐nonyl butyrate. The importance of absolute configuration on attractiveness, the effects on trap catches of the addition of minor pheromone components, dispenser type, and pheromone loading are described in the development of an optimised pheromone lure with which to trap H. marginata males. In analyses of volatiles collected from virgin female H. marginata by gas chromatography (GC) coupled with electroantennographic recording (EAG) from the antenna of a male H. marginata, two EAG responses were observed. Analyses by coupled GC‐mass spectrometry (MS) indicated these were due to 2‐nonyl butyrate and a trace amount (1%) of 2‐heptyl butyrate. A similar trace amount of 2‐nonanol was detected in GC‐MS analyses but this compound did not elicit an EAG response when the synthetic compound was tested, whereas the other two compounds did. These three compounds were not observed in collections of volatiles made from male H. marginata. The 2‐nonyl butyrate was shown to be the (R)‐enantiomer. In field trapping tests (R)‐2‐nonyl butyrate was at least 10× more attractive to male H. marginata than the racemic compound, and the (S)‐enantiomer was unattractive. Addition of the potential minor components individually or together at the naturally occurring ratios did not increase or reduce the attractiveness of the lure. Polyethylene vials and rubber septa were equally effective as pheromone dispensers, lasting for at least 5 weeks in the field in the UK, although laboratory tests indicated release from the former was more uniform and more likely to last longer in the field. Increasing loading of pheromone in the dispenser increased attractiveness. Traps baited with polyethylene vials containing 0.5 mg of (R)‐2‐nonyl butyrate are recommended for monitoring H. marginata and these are far more sensitive than water or sticky traps currently used for monitoring this pest.

Factors affecting trap catch in pheromone-based monitoring of saddle gall midge Haplodiplosis marginata (Diptera: Cecidomyiidae)

BACKGROUND The saddle gall midge, Haplodiplosis marginata (von Roser) (Diptera: Cecidomyiidae), is a pest of cereal crops in Europe. Outbreaks are difficult to predict and effective monitoring tools are required to ensure the effectiveness of pest management options. The female sex pheromone (R)-2-nonyl butyrate provides the basis of a highly effective lure for this insect. Here, we demonstrate how the success of this lure can be influenced by parameters such as trap location, lure age, and interference between traps fitted with these lures. RESULTS A pheromone lure containing (R)-2-nonyl butyrate attracted male midges for at least 9 weeks under field conditions. Pheromone-baited traps performed best when situated away from field margins and below the height of the crop. Interference between nearby traps was evident at distances <20 m. CONCLUSION The results presented here offer new insights into the behavioural responses of male H. marginata to the female sex pheromone and provide practical recommendations for the use of H. marginata pheromone traps in the field.

Recording the movement of adult vine weevil within strawberry crops using radio frequency identification tags

BACKGROUND:Vine weevil (Otiorhynchus sulcatus) is a major pest of soft fruit and ornamental crops. There is an urgent need to improve control of vine weevil and in particular to provide growers with effective Integrated Pest Management-compatible controls with which to target the adult stage of this pest. One approach would be to exploit the behaviour of adult vine weevil to disseminate spores of an entomopathogenic fungus placed within the crop environment in artificial refuges. To be effective this approach requires that the weevils move through the crop environment and in doing so spread the pathogen from the artificial refuges. OBJECTIVE:Use passive radio frequency identification (RFID) tags to study the movement of adult vine weevil within crop environments. METHOD:A series of laboratory bioassays were completed in which the effect of attaching RFID tags using a thermoplastic or a cyanoacrylate adhesive on survival and movement, on both horizontal and vertical surfaces, of adult vine weevil was determined. An outdoor field experiment was then completed at Harper Adams University in order to test the potential of this technique for studying vine weevil movement within crop environments. RESULTS:Attaching RFID tags using the thermoplastic adhesive did not result in any weevil deaths over a 21 day period. In contrast, just over half (53%) of the weevils to which the RFID tag was attached using the cyanoacrylate adhesive died over the same period. The mean of weevil horizontal movement speed was significantly slower when an RFID tag was attached using a thermoplastic (1.01 cm/s) or a cyanoacrylate (0.29 cm/s) adhesive compared with untagged weevils (1.83 cm/s). However, weevils that were tagged using the thermoplastic adhesive were significantly faster than weevils tagged using the cyanoacrylate adhesive. Mean vertical movement speed was also significantly slower when weevils were tagged using the thermoplastic adhesive (0.18 cm/s) compared with untagged weevils (0.37 cm/s). Weevils tagged using the cyanoacrylate adhesive were unable to climb vertical surfaces.In the field experiment, weevils moved away from their release points. Nine days after the start of the experiment weevils were on average 3.38 m from their release points indicating a speed of movement of 0.38 m/day. The mean distance of movement from their release points did not increase further during the rest of the experimental period, but remained relatively constant at between 2.50 and 3.28 m. As such, for weevils that remained within the crop environment, there is no evidence of dispersal behaviour,with movement behaviour observed more likely to be driven by resource utilisation. However, not all weevils remained within the crop area. Indeed, 15 (38%) of released weevils and/or RFID tags left the crop area, indicating possible long range dispersal by these individuals or evidence of predation of the weevils.A total of 11 (28%) of the RFID tagged weevils released into the crop were recovered alive and with the tag still attached after 35 days. These weevils were estimated to have moved a distances of between 2.65 and 17.30 m (average distance moved 7.50 m) during this period. These distances are likely to underestimate the distance moved by each weevil as they assume that each weevil took the most direct route between each point and did not move other than this. In total eight of the RFID-tagged weevils moved both along and between rows of strawberry grow-bags. At the start of the experiment the 11 RFID tagged weevils occupied 11 (14%) of the strawberry grow-bags. If these weevils took the most direct route between each position within the crop where they were detected these weevils would have crossed, and potentially laid eggs in, 44 (58%) of the grow-bags during the 35 days of the experiment. If they had taken a more indirect route the weevils could have potentially laid eggs in a higher number of the grow-bags. CONCLUSIONS:Results presented here indicate that RFID tags can be used to study the movement of vine weevil adults within crop environments. However, the weight and size of currently available tags significantly slows the movement of weevils under laboratory conditions and frequency of detection may affect estimates of actual distance moved. Despite this, the rate at which vine weevil dispersed through the strawberry crop was comparable to the speed of movement recorded previously by others when weevils were released into an urban environment. Use of RFID tags also resulted in detection rates far higher than those reported in studies by others using traditional mark-release-recapture techniques. Use of RFID tags in the present study indicates that adult vine weevil have the potential to disperse spores of a suitable entomopathogenic fungus from artificial refuges throughout the crop environment. Use of this technique could also be applied to investigate the effect of other plant protection products as well as the impact of different cropping systems on vine weevil movement and survival.