Effective early detection, monitoring and management methods are critical for reducing the impacts of insect pests in agriculture and forestry. Combining host plant volatiles with sex pheromones could enhance trapping methodologies, whilst the use of non-host volatiles could improve the effectiveness of pest management through repellency effects. In this meta-analysis approach, we analysed 51 studies that used electroantennograms (EAG), wind tunnels and/or field traps to evaluate the antennal and behavioural responses of Lepidoptera to sex pheromones combined with attractant or repellent plant volatiles. Proposed attractant plant volatiles had a positive association with female Lepidoptera responses to sex pheromone, but effects on males were highly variable, with unexpected repellency reported in some studies. Proposed repellent plant volatiles were significantly or near-significantly negatively associated with male attraction to sex pheromones but were scarcely studied. Sub-group analysis identified that male responses to sex pheromone were reduced when the dose of attractant plant volatile relative to sex pheromone was increased. Green-leaf volatiles were associated with the strongest positive effects for males in field traps. Multiple-compound attractant plant volatile blends were less effective than single compounds in field studies. Our analysis demonstrates, (i) the potential value of combining host plant volatiles with sex pheromones to capture females rather than only males, (ii) the importance of identifying appropriate host plant volatiles and optimal relative doses, and (iii) the potential for non-host plant volatile use in pest management strategies.

Methyl isonicotinate is one of several patented 4-pyridyl carbonyl compounds being investigated for a variety of uses in thrips pest management. It is probably the most extensively studied thrips non-pheromone semiochemical, with field and glasshouse trapping experiments, and wind tunnel and Y-tube olfactometer studies in several countries demonstrating a behavioural response that results in increased trap capture of at least 12 thrips species, including the cosmopolitan virus vectors such as western flower thrips and onion thrips. Methyl isonicotinate has several of the characteristics that are required for an effective semiochemical tool and is being mainly used as a lure in combination with coloured sticky traps for enhanced monitoring of thrips in greenhouses. Research indicates that this non-pheromone semiochemical has the potential to be used for other thrips management strategies such as mass trapping, lure and kill, lure and infect, and as a behavioural synergist in conjunction with insecticides, in a range of indoor and outdoor crops.

The rice stem bug, Tibraca limbativentris Stal. (Hemiptera: Pentatomidae) is one of the most important pests of rice crops, especially irrigated crops. Plant defence strategies against these bugs may involve the emission of chemical compounds, which are released following herbivore attacks, directly or indirectly harming pest performance. The aim of this study was to evaluate the influence of constitutive and herbivory-induced volatiles from rice plants (Oryza sativa L.) on the behavioural responses of T. limbativentris adults and egg parasitoids Trissolcus basalis (Wollaston) and Telenomus podisi (Ashmead) (Hymenoptera, Platygastridae). Plant volatiles were collected from undamaged plants of the rice cultivar IRGA 424 and from plants that suffered herbivory by five males or five females of T. limbativentris. Air-entrainment extracts were analysed by GC–flame ionization detector and GC–MS, and insect responses evaluated in a ‘Y’ olfactometer. T. limbativentris feeding damaged on rice plants induced the release of 16 volatiles compounds in a higher amounts compared to undamaged plants The main compounds induced were (E)-2-hexenal, (E)-2-octen-1-ol, methyl salicylate and α-muurolene. Female bugs were significantly attracted to air-entrainment extracts containing volatiles from undamaged plants compared with air-entrainment extracts containing volatiles emitted from plants damaged by T. limbativentris, whereas males showed no preference. Telenomus podisi females were significantly attracted to volatiles from air-entrainment extracts of plants damaged by females, whereas T. basalis showed no preference. These results suggest that rice plants may be emitting defence compounds, which could be avoided by T. limbativentris females and also acted indirectly by attracting natural enemies.

Wheat bulb fly (WBF) larvae use chemotaxis to orientate towards host-plant root exudates. This study aimed to investigate the role of the primary plant metabolite carbon dioxide (CO2) in host-plant location by WBF. Arena based behavioural experiments were used to identify whether CO2 induced chemotaxis (directional movement in response to a chemical stimulus) or kinesis (non-directional movement in response to a stimulus) from WBF larvae. No chemotactic response was observed when larvae were presented to a point source of CO2. However, elevated levels of CO2 induced kinesis, with both track length and tortuosity (number of twists and turns in the movement path) increasing at elevated CO2 levels of 1000–2000 ppm, demonstrating increased searching behaviour. Soil emission of CO2 was quantified to compare soil levels with those identified as eliciting behavioural effects on the larvae. Samples removed from soil gave a mean CO2 concentration of 557 (±46) ppm, which is lower than the lowest concentration of CO2 found to induce a behavioural response and higher than the lowest CO2 concentration tested, which was found not to alter behaviour. It is proposed that increased CO2 concentrations in the soil act as a behavioural trigger, inducing intensive searching of an area by WBF larvae. This increases the likelihood of finding more host-specific identifiers, such as secondary metabolites when near a potential host-plant.

The aphid sex pheromone component (4aS,7S,7aR)-nepetalactone is considered to be a potential tool for enhancing biological control of aphids. Studies have confirmed its potential to attract parasitoids, increase parasitism rates in the field and also alter the spatial distribution of parasitoids. An important aspect that has been overlooked is the impact that the introduction of nepetalactone may have on aphid populations already present in field or glasshouse environments. The most prevalent pest aphid populations in glasshouse and field environments are the asexual morphs, which are capable of exponential growth if populations are not controlled. The short-term implications of the sex pheromone on asexual aphids were observed through their behavioural response. Using Y-tube olfactometry, it is shown that virginoparae of the peach-potato aphid, Myzus persicae, are repelled by high concentrations of nepetalactone. Long-term effects of the pheromone which may span the aphid's life, or even generations, were assessed via mean relative growth rate (MRGR) and the intrinsic rate of natural increase (rm). Electroantennography also demonstrated that asexual female aphids are able to detect aphid sex pheromone components. To our knowledge, this is the first time it has been reported that M. persicae virginoparae are able to detect aphid sex pheromone components or that their behavioural response and/or performance has been studied. The implications of these results and their significance in understanding semiochemical communication are discussed.

The life cycle of the parasitic copepod Lernaeocera branchialis involves 2 hosts, typically a pleuronectiform host upon which development of larvae and mating of adults occurs and a subsequent gadoid host, upon which the adult female feeds and reproduces. Both the copepodid and adult female stages must therefore locate and identify a suitable host to continue the life cycle. Several mechanisms are potentially involved in locating a host and ensuring its suitability for infection. These may include mechano-reception to detect host movement and chemo-reception to recognize host-associated chemical cues, or kairomones. The aim of this study was to identify the role of kairomones in host location by adult L. branchialis, by analysing their behaviour in response to fish-derived chemicals. Experiments demonstrated that water conditioned by immersion of whiting, Merlangius merlangus, elicited host-seeking behaviour in L. branchialis, whereas cod- (Gadus morhua) conditioned water did not. Lernaeocera branchialis are considered a genetically homogeneous population infecting a range of gadoids. However, their differential response to whiting- and cod-derived chemicals in this study suggests that either there are genetically determined subspecies of L. branchialis or there is some form of environmental pre-conditioning that allows the parasite to preferentially recognize the host species from which it originated.

Wheat bulb fly (WBF, Delia coarctata, Fallén, Diptera: Anthomyiidae) is a pest of commercial importance in wheat, barley and rye, with attacked crops failing to produce full potential yields. Females do not oviposit in association with a host-plant; therefore, prompt location of a suitable host is critical to the survival of the newly hatched larvae. The objective of this study was to conduct choice test bioassays to assess the attraction of WBF larvae to specific chemical constituents of WBF host-plant root exudates, the hydroxamic acids DIMBOA (2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one) and MBOA (6-methoxy-benzoxazolin-2-one). The larval response to four concentrations of each test compound was assessed in arena bioassays. Analysis using a Rayleigh test of uniformity of the final resting positions of larvae in response to these chemicals indicated attraction. These results go some way to explaining the mechanisms by which WBF larvae locate host plants, giving the potential to develop semiochemical based control strategies.

The common bed bug, Cimex lectularius (Hemiptera: Cimicidae), has recently re-emerged in increasing numbers, distribution and intensity of infestation in many countries. Current control relies on the application of residual pesticides; but, due to the development of insecticide resistance, there is a need for new tools and techniques. Semiochemicals (behaviour and physiology modifying chemicals) could be exploited for management of bed bugs. However, in order to identify semiochemicals that can be utilised in monitoring or control, a suitable olfactometer is needed that enables the study of the responses of bed bugs to volatile chemicals. Previous studies have used olfactometers that do not separate olfactory responses from responses to physical contact. In this study, a still-air olfactometer was used to measure behavioural responses to different bed bug-derived volatiles presented in an odour pot. Bed bugs were significantly more likely to visit the area above the odour pot first, and more frequently, in the presence of volatiles from bed bug-exposed paper but not in the presence of volatiles from conspecific bed bugs. Bed bug activity was found to be dependent on the presence of the volatiles from bed bug-exposed paper, the time during the scotophase and the sex of the insect being tested. The still-air olfactometer could be used to test putative semiochemicals, which would allow an understanding of their behavioural role in bed bug ecology. Ultimately, this could lead to the identification of new semiochemical tools for bed bug monitoring and control.

Mosquitoes such as Culex quinquefasciatus Say (Diptera: Culicidae) are important vectors of organisms that cause disease in humans. Research into the development of effective standardized odour baits for blood-fed females (oviposition attractants), to enable entomological monitoring of vector populations, is hampered by complex protocols for extraction of physiologically active volatile chemicals from natural breeding site water samples, which have produced inconsistent results. Air entrainment and solvent extraction are technically demanding methods and are impractical for use in resource poor environments where mosquito-borne disease is most prevalent. This study reports the first use of a simple, robust extraction technique, stir bar sorptive extraction (SBSE), to extract behaviourally active small lipophilic molecules (SLMs) present in water samples collected from Cx. quinquefasciatus breeding sites in Tanzania. Extracts from a pit latrine and from a cess pool breeding site attracted more gravid Cx. quinquefasciatus in pair choice bioassays than control extracts, and coupled gas chromatography-electroantennography (GC-EAG) allowed tentative identification of 15 electrophysiologically active chemicals, including the known oviposition attractant, skatole (3-methylindole). Here, we have demonstrated, using simple pair choice bioassays in controlled laboratory conditions, that SBSE is effective for the extraction of behaviourally and electrophysiologically active semiochemicals from mosquito breeding site waters. Further research is required to confirm that SBSE is an appropriate technique for use in field surveys in the search for oviposition cues for Cx. quinquefasciatus.

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.

Selected studies in chemical ecology are described that have led to the identification of semiochemicals playing key roles in pest behaviour. In some instances, these compounds can provide alternatives to insecticides, but novel integrated control programmes including use of semiochemicals are described which have greater potential for the future. Thus, semiochemicals can be used to increase the efficacy of entomophagous fungi and to manipulate populations of beneficial insects. New strategies are being developed which use both antifeedant and attractant semiochemicals to deter insects from crops and aggregate them in areas where they can be destroyed. For the long term, strategies are described that employ genetic manipulation of crop plants so that pests are controlled by semiochemicals produced directly in the plants.