Behavior-modifying Chemicals Research Articles

D-Limonene Is the Active Olfactory Attractant in Orange Juice for Bactrocera dorsalis (Insecta: Diptera: Tephritidae).

The oriental fruit fly, Bactrocera dorsalis (Hendel), poses a significant threat to the global fruit industry, causing damage to diverse fruits like citrus, mango, and guava. Chemical pesticides have limited effectiveness, and pesticide residues and pesticide resistance are pressing issues. Therefore, it is essential to develop environmentally friendly pest control methods to address this problem. Behavior-modifying chemicals, including male attractants and intersex protein baits, play a critical role in the control of B. dorsalis. The mature host fruit serves as both an oviposition site and food source under natural conditions, making it a potential attraction source for oriental fruit flies. Orange, Citrus sinensis, is a main host of B. dorsalis, and commercial orange juice is a common attractant for the egg laying of B. dorsalis. Although it can both attract and elicit oviposition behaviors in B. dorsalis adults, its active components are still unclear. This study utilized analytical chemistry, behavioral tests, and electrophysiology to identify the active components of commercial orange juice that attract B. dorsalis, with the aim of providing a reference for the development of behavior-modifying chemical-based techniques to control B. dorsalis. Five compounds with a high abundance were identified via a GC-MS, including D-Limonene, butanoic acid ethyl ester, β-myrcene, linalool, and α-terpineol. Behavioral and electrophysiological experiments uncovered that D-Limonene was the active substance that was the main attractant in the mixture of these five substances, evoking a strong electrophysiological response in adult B. dorsalis. D-Limonene strongly attracts adult B. dorsalis only when they are sexually mature, and the attraction is not rhythmic. Olfaction plays a leading role in the attraction of D-Limonene to adult B. dorsalis, and Orco-/- mediates the perception of D-Limonene by B. dorsalis. Overall, D-Limonene is one of the key attractant compounds for B. dorsalis in the volatile compounds of commercial orange juice, offering possible support for the development of behavior-modifying chemical-based technology to control B. dorsalis in the future.

A classic screening marker does not affect antennal electrophysiology but strongly regulates reproductive behaviours in Bactrocera dorsalis.

The key phenotype white eye (white) has been used for decades to selectively remove females before release in sterile insect technique programs and as an effective screening marker in genetic engineering. Bactrocera dorsalis is a representative tephritid pest causing damage to more than 150 fruit crops. Yet, the function of white in important biological processes remains unclear in B. dorsalis. In this study, the impacts of the white gene on electrophysiology and reproductive behaviour in B. dorsalis were tested. The results indicated that knocking out Bdwhite disrupted eye pigmentation in adults, consistent with previous reports. Bdwhite did not affect the antennal electrophysiology response to 63 chemical components with various structures. However, reproductive behaviours in both males and females were significantly reduced in Bdwhite-/- . Both pre-copulatory and copulation behaviours were significantly reduced in Bdwhite-/- , and the effect was male-specific. Mutant females significantly delayed their oviposition towards γ-octalactone, and the peak of oviposition behaviour towards orange juice was lost. These results show that Bdwhite might not be an ideal screening marker in functional gene research aiming to identify molecular targets for behaviour-modifying chemicals. Instead, owing to its strong effect on B. dorsalis sexual behaviours, the downstream genes regulated by Bdwhite or the genes from white-linked areas could be alternate molecular targets that promote the development of better behavioural modifying chemical-based pest management techniques.

Sensing of Airborne Infochemicals for Green Pest Management: What Is the Challenge?

One of the biggest global challenges for our societies is to provide natural resources to the rapidly expanding population while maintaining sustainable and ecologically friendly products. The increasing public concern about toxic insecticides has resulted in the rapid development of alternative techniques based on natural infochemicals (ICs). ICs (e.g., pheromones, allelochemicals, volatile organic compounds) are secondary metabolites produced by plants and animals and used as information vectors governing their interactions. Such chemical language is the primary focus of chemical ecology, where behavior-modifying chemicals are used as tools for green pest management. The success of ecological programs highly depends on several factors, including the amount of ICs that enclose the crop, the range of their diffusion, and the uniformity of their application, which makes precise detection and quantification of ICs essential for efficient and profitable pest control. However, the sensing of such molecules remains challenging, and the number of devices able to detect ICs in air is so far limited. In this review, we will present the advances in sensing of ICs including biochemical sensors mimicking the olfactory system, chemical sensors, and sensor arrays (e-noses). We will also present several mathematical models used in integrated pest management to describe how ICs diffuse in the ambient air and how the structure of the odor plume affects the pest dynamics.

Alternative Methods of Ant (Hymenoptera: Formicidae) Control with Emphasis on the Argentine Ant, Linepithema humile.

Simple SummaryAnts can be major pests to homeowners and other property owners. In the U.S., ants often rank as one of the most common and difficult-to-control pests around and in homes, businesses, and other facilities. Typically, ant control practices are conducted by licensed pest management professionals with sprays, baits, and granular products, containing various types of insecticides, applied to the outside perimeter of infested structures. Many of the insecticides used to control pest ants are harmful to non-target organisms, especially those in aquatic environments. To address these negative environmental impacts, research on alternative, generally low-impact and least toxic, ant control strategies has received a fair amount of attention. The underlying goal of this research is a reduction in human exposure to traditional insecticides. Examples of alternative approaches included in this review article include the use of essential oils and other chemicals as deterrents to ant nesting and foraging; ant trail pheromones as disruptants to foraging; mass trapping of ants; new gel baits containing extremely low concentrations of insecticide; and altering the behavior and distribution of ants by altering their access to food.Ants (Hymenoptera: Formicidae), especially the Argentine ant, Linepithema humile (Mayr), can be significant nuisance pests in urban and suburban environments. Conventional interventions have primarily relied on the use of chemical insecticides, namely fipronil and bifenthrin, applied as residual, contact treatments around the outside perimeter of infested structures. Despite tightening regulation limiting the scope of insecticide applications in urban settings, dependence on these products to manage ants continues, resulting in significant water contamination. The U.S. EPA, in response, has further restricted the use patterns of many insecticides used for ant control in professional and over-the-counter markets. The purpose of this review is to summarize the relevant literature associated with controlling nuisance pest ants, with emphasis on L. humile, without the use of liquid broadcast applications of EPA-registered insecticides while focusing on low-impact, alternative (to broadcast applications) pest control methods. Specific subsections include Trail Pheromone; Use of Behavior-Modifying Chemicals; Mass Trapping; Hydrogels, “Virtual” Baiting, and Exceedingly-Low Bait Concentrations; Food Source Reduction; Deterrents; and RNA Interference (RNAi).

13-Tetradecenyl acetate, a female-produced sex pheromone component of the economically important click beetle Melanotus communis (Gyllenhal) (Coleoptera: Elateridae)

Species-specific behavior-modifying chemicals have been used for more than 50 years for monitoring and management of insect pests of agriculture and human health. Elaterid beetle larvae are among insect pests in soil that are increasingly problematic, in part due to the lack of effective management strategies. However, little is known about the insect-produced chemicals that mediate the reproductive behavior of these pests. We used chemical and behavioral studies to identify, synthesize, and field test the sex attractant pheromone of adults of Melanotus communis, commonly called the corn wireworm, the larvae of which are economically important pests of U.S. crops. Our results indicated that a single female-produced chemical, 13-tetradecenyl acetate, was strongly attractive to conspecific male beetles, and did not appear to attract other species. In field evaluations, male M. communis exhibited a dose-dependent response to this compound. In a trial comparing different slow-release dispensers, a small rubber septum impregnated with the chemical was as effective as and easier to use than a plastic bag dispenser. Given that the sex attractant of this insect consists of a single compound that can be readily synthesized, its development for monitoring and management of the corn wireworm may be economically feasible.

A Multiple-Choice Bioassay Approach for Rapid Screening of Key Attractant Volatiles.

Fermentation volatiles attract a wide variety of insects and are used for integrated pest management. However, identification of the key behavior modifying chemicals has often been challenging due to the time consuming nature of thorough behavioral tests and unexpected discrepancies between laboratory and field results. Thus we report on a multiple-choice bioassay approach that may expedite the process of identifying field-worthy attractants in the laboratory. We revisited the four-component key chemical blend (acetic acid, ethanol, acetoin, and methionol) identified from 12 antennally active wine and vinegar chemicals for Drosophila suzukii (Matsumura) (Diptera: Drosophilidae). The identification of this blend took 2 yr of continuous laboratory two-choice assays and then similarly designed field trials. This delay was mainly due to a discrepancy between laboratory and field results that laboratory two-choice assay failed to identify methionol as an attractant component. Using a multiple-choice approach, we compared the co-attractiveness of the 12 potential attractants to an acetic acid plus ethanol mixture, known as the basal attractant for D. suzukii, and found similar results as the previous field trials. Only two compounds, acetoin and, importantly, methionol, increased attraction to a mixture of acetic acid and ethanol, suggesting the identification of the four-component blend could have been expedited. Interestingly, the co-attractiveness of some of the 12 individual compounds, including a key attractant, methionol, appears to change when they were tested under different background odor environments, suggesting that background odor can influence detection of potential attractants. Our findings provide a potentially useful approach to efficiently identify behaviorally bioactive fermentation chemicals.

English

Behaviour-modifying chemicals like pheromones and kairomones hold a great potential in pest management. Evidences from mating behaviour studies of the banana weevil, and from the weevil’s responses to their freeze-killed conspecifics, body washes/extracts, live conspecifics (olfactometer studies), and trapped volatiles of mature and immature adults clearly suggest that two types of pheromones are produced in this insect: a female produced sex pheromone and a male produced aggregation pheromone. Both are perceived by olfactory means. The latter has already been isolated by earlier workers and is in use in control programs. Greater successes may however, be recorded with the control of this pest (e.g. in mating disruptions, mass trappings, pest monitoring) if the female-sex pheromone also gets finally isolated, and used in conjunction with good cultural practices. Key words: Cosmopolites sordidus, sex pheromone, mating disruptions, bioassays, olfactometer studies, gas chromatographic profiles, electroantennogram studies.

Olfactory responses of western flower thrips (Frankliniella occidentalis) populations to a non-pheromone lure

Western flower thrips (WFT), Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), is a major pest of horticultural crops worldwide. The search for alternative pest management techniques has resulted in increasing interest in the use of kairomones and other behaviour-modifying chemicals to mitigate the impact of this pest. In this study, we determined whether the origin of populations, feeding history, and/or genotype influence the response of WFT to the thrips kairomone lure methyl isonicotinate (MI) in a Y-tube olfactometer study. Four New Zealand thrips populations were tested: (1) from a commercial glasshouse capsicum crop, (2) from a long-established laboratory colony (>222 generations) kept on chrysanthemums, (3) from a laboratory colony (6–9 generations) kept on French dwarf beans, and (4) thought to be a separate cryptic non-pest species from outdoor yellow tree lupins, Lupinus arboreus Sims (Fabaceae). In the laboratory tests, significantly more WFT from all four populations chose the MI-laden arm of a Y-tube olfactometer when it contained 1 μl methyl isonicotinate (61.3–73.2%) compared with the blank no-odour arm. No differences in response to MI were found between the two laboratory and the one glasshouse WFT populations. Both laboratory populations and the greenhouse population belonged to the ‘glasshouse pest’ genotype of WFT. However, the cryptic non-pest WFT genotype responded more strongly to MI than any of the other populations, although the response was only significantly stronger than that of the long-established laboratory population. Significant differences were also found among populations in the average time taken for thrips to make a choice to enter either arm of the Y-tube olfactometer, with the cryptic non-pest lupin genotype taking the shortest time, followed by thrips from the capsicum glasshouse. The results are discussed with respect to the variability in olfactory perception and olfactory behaviour within a species and the relevance to the use of such a kairomone lure in pest management programmes.

Novel high-throughput screens of Anopheles gambiae odorant receptors reveal candidate behaviour-modifying chemicals for mosquitoes.

Despite many decades of multilateral global efforts, a significant portion of the world population continues to be plagued with one or more mosquito-vectored diseases. These include malaria and filariasis as well as numerous arboviral-associated illnesses including Dengue and Yellow fevers. The dynamics of disease transmission by mosquitoes is complex, and involves both vector competence and vectorial capacity. One area of intensive effort is the study of chemosensory-driven behaviours in the malaria vector mosquito Anopheles gambiae Giles, the modulation of which are likely to provide opportunities for disease reduction. In this context recent studies have characterized a large divergent family of An. gambiae odorant receptors (AgORs) that play critical roles in olfactory signal transduction. This work has facilitated high-throughput, cell-based calcium mobilization screens of AgOR-expressing HEK cells that have identified a large number of conventional AgOR ligands, as well as the first non-conventional Orco (olfactory receptor co-receptor) family agonist. As such, ligand-mediated modulation serves as a proof-of-concept demonstration that AgORs represent viable targets for high-throughput screening and for the eventual development of behaviour-modifying olfactory compounds. Such attractants or repellents could foster malaria reduction programmes.

Age and Sex Related Responsiveness ofTribolium castaneum(Coleoptera: Tenebrionidae) in Novel Behavioral Bioassays

The hardiness and mobile nature of Tribolium castaneum (Herbst) make them easy to work with but are the same factors that make their responses to behavior-modifying chemical stimuli difficult to evaluate. To overcome these difficulties two bioassays were developed: a two-choice test with airflow and a diffusion-based test to evaluate responses to chemical stimuli. The two-choice assay is excellent for rapidly comparing two stimuli or examining the response to one stimulus against a control. The diffusion assay determines differences in orientation behavior to multiple simultaneous stimuli and can examine other behaviors during exposure. Preparation of individuals for bioassay is also important, because disturbance increases the activity level of individual beetles beyond the duration of the disturbance. The age and the sex of beetles affect responsiveness to chemical cues. These bioassays and a better understanding of T. castaneum's activity have revealed approaches for evaluating its responsiveness to behavior-modifying chemicals.

Formate Analogs as Antagonists of the Sex Pheromone of the Honeydew Moth, Cryptoblabes gnidiella: Electrophysiological, Behavioral and Field Evidence

Cryptoblabes gnidiella Milliére (Lepidoptera: Pyralidae) is an economically important exotic pest of vineyards in Southern Brazil and Uruguay. The sex pheromone of C. gnidiella was identified as a mixture of (Z)-11-hexadecenal and (Z)-13-octadecenal, and has been used to monitor populations of this pest in Israel. The development of mating disruption for this species may be hampered by the chemical instability of the natural pheromone components. Therefore, studies on more stable pheromone analogs may provide tools for a control strategy based on behavior-modifying chemicals. We report here the electrophysiological and behavioral responses of C. gnidiella males to (Z)-9-tetradecenyl formate and (Z)-11-hexadecenyl formate, structural analogs of the pheromone components. In gas chromatography-electroantennogram detection (GC-EAD) studies, both analogs elicited responses from C. gnidiella male antennae. Pre-exposure to the formates did not affect the subsequent EAD responses to the natural pheromone components. The formates acted as pheromone antagonists in wind tunnel tests, inhibiting the responses of males toward both synthetic pheromone and calling females. In the field, captures of males in pheromone-baited traps decreased, in a dose-response pattern, when different amount of formates were added to the pheromone. These pheromone antagonists, thus, are potentially useful as mating disruptants for C. gnidiella in commercial vineyards.

A bioassay for studying behavioural responses of the common bed bug, Cimex lectularius (Hemiptera: Cimicidae) to bed bug-derived volatiles

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.

Sex Pheromones and Their Impact on Pest Management

The idea of using species-specific behavior-modifying chemicals for the management of noxious insects in agriculture, horticulture, forestry, stored products, and for insect vectors of diseases has been a driving ambition through five decades of pheromone research. Hundreds of pheromones and other semiochemicals have been discovered that are used to monitor the presence and abundance of insects and to protect plants and animals against insects. The estimated annual production of lures for monitoring and mass trapping is on the order of tens of millions, covering at least 10 million hectares. Insect populations are controlled by air permeation and attract-and-kill techniques on at least 1 million hectares. Here, we review the most important and widespread practical applications. Pheromones are increasingly efficient at low population densities, they do not adversely affect natural enemies, and they can, therefore, bring about a long-term reduction in insect populations that cannot be accomplished with conventional insecticides. A changing climate with higher growing season temperatures and altered rainfall patterns makes control of native and invasive insects an increasingly urgent challenge. Intensified insecticide use will not provide a solution, but pheromones and other semiochemicals instead can be implemented for sustainable area-wide management and will thus improve food security for a growing population. Given the scale of the challenges we face to mitigate the impacts of climate change, the time is right to intensify goal-oriented interdisciplinary research on semiochemicals, involving chemists, entomologists, and plant protection experts, in order to provide the urgently needed, and cost-effective technical solutions for sustainable insect management worldwide.

The influence of fluorinated molecules (semiochemicals and enzyme substrate analogues) on the insect communication system.

Can the introduction of fluorine atoms affect the bioactivity of natural semiochemicals? Can fluorine contribute in the creation of specific enzyme inhibitors to interrupt or disrupt the insect communication system? The first step for the bioactivity of a molecule is interaction with the biological sensor. Hydrogen and fluorine are almost bioisosteric and the receptor site of the enzyme can still recognize and accept the fluoro analogue of its natural substrate. However, the peculiar electronegativity of the fluorine atom can affect the binding, absorption, and transport of the molecule. The differences in the molecule's electronic properties can lead to differences in the chemical interactions between the receptor and the fluorinated substrate. Fluorine introduction can modify the metabolic stability and pathway of the semiochemicals in many different ways. Fluorinated analogues can show synergism, inhibition, or hyperagonism effects on insect behaviors, that is, the activity of the nonfluorinated parent compounds can be mimicked, lost, or increased. In any case, the fluorinated molecules can interact with the bioreceptors in a new and disrupting way. The semiochemicals are olfactory substances: fluorine can affect their volatility or smell. Production of semiochemicals from exogenous substances, perception at antennal receptors, and processing of biological responses are the main steps of communication among insects. In the production step, the fluorinated molecules can interact with enzymes that catalyze the biosynthesis of the natural pheromones. In the perception step, fluorinated semiochemicals can interact with the olfactory receptor cells; this often leads to totally unpredictable behaviors. Fluorinated molecules have been developed as probes to elucidate the complex chemorecognition processes of insects. Many of these molecules have been tested to find highly effective behavior-modifying chemicals. New analogues have been synthesized to investigate the metabolic pathway of a pheromone molecule and many of them are promising disrupting agents. Despite such titanic research efforts, the results have often been random, rational trends in the induced behaviors have sometimes been impossible to find, and practical applications of the fluorinated semiochemicals are still uncertain.

Development of field strategies incorporating semiochemicals for the control of the pea and bean weevil, Sitona lineatus L.

The use of semiochemicals (behaviour-modifying chemicals) as components of a control strategy for Sitona lineatus (Coleoptera: Curculionidae), was investigated in two field trials. The semiochemicals used were 4-methyl -3,5-heptanedione, the aggregation pheromone of S. lineatus, and an antifeedant formulation of neem oil, from the neem tree, Azadirachta indica (Meliaceae). The results showed that, in small-plot trials, insect damage to the crop could be decreased by the application of an antifeedant. Plots baited with pheromone had more adult and larval damage than untreated plots, indicating increased insect activity. This work demonstrated that in principle, populations of S. lineatus can be manipulated with semiochemicals when these arc deployed in a ‘push-pull’ or ‘stimulo-deterrent diversionary’ strategy. Such a strategy, in which insects are diverted from the crop by a feeding deterrent and attracted to discard areas with an attractant pheromone, could be developed for the management of this pest in arable agriculture.

Insect pheromones and other behaviour-modifying chemicals: applications and regulation. Edited by R.L. Ridgway, May Inscoe & H. Arn. BCPC Monograph No. 51. (Farnham, Surrey: The British Crop Protection Council, 1992). x + 135 pp. Soft cover £15.00. ISBN 0-948404-62-0.

Insect pheromones and other behaviour-modifying chemicals: applications and regulation. Edited by R.L. Ridgway, May Inscoe & H. Arn. BCPC Monograph No. 51. (Farnham, Surrey: The British Crop Protection Council, 1992). x + 135 pp. Soft cover £15.00. ISBN 0-948404-62-0.

Behavior‐Modifying Chemicals for Insect Management. Applications of Pheromones and Other Attractants

Journal of Environmental QualityVolume 20, Issue 1 p. 313-313 Book Review Behavior-Modifying Chemicals for Insect Management. Applications of Pheromones and Other Attractants Edited by Richard L. Ridgway, Robert M. Silverstein, and May N. Inscoe, Marcel Dekker, Inc., 270 Madison Avenue, New York, NY 10016. 1990. 761 p. $195.00 (domestic), $234.00 (abroad). ISBN 0-8247-8156-2 R.A.J. Taylor, R.A.J. Taylor Department of Entomology, Ohio Agricultural and Development Center, Wooster, OH, 44691Search for more papers by this author R.A.J. Taylor, R.A.J. Taylor Department of Entomology, Ohio Agricultural and Development Center, Wooster, OH, 44691Search for more papers by this author First published: 01 January 1991 https://doi.org/10.2134/jeq1991.00472425002000010059xCitations: 2AboutPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinkedInRedditWechat No abstract is available for this article.Citing Literature Volume20, Issue1January‐March 1991Pages 313-313 RelatedInformation

Behavior-Modifying Chemicals for Insect Management: Applications of Pheromones and Other Attractants. Based on a Symposium. Richard L. Ridgway , Robert M. Silverstein , May N. Inscoe

<i>Behavior-Modifying Chemicals for Insect Management: Applications of Pheromones and Other Attractants. Based on a Symposium</i>. Richard L. Ridgway , Robert M. Silverstein , May N. Inscoe

Review: Behavior-modifying chemicals for insect management

Ridgway, R. L., Silverstein, R. M. &amp; Inscoe, M. N. (eds.) 1990: Behavior-modifying chemicals for insect management. Applications of pheromones and other attractants. - Marcel Dekker Inc., New York, USA. 761 pp.ISBN 0-8247 -8156-2. USD 195 in the USA and Canada, and USD 234 elsewhere.

Ecology and management of forest insects

Part 1 Introduction: forests as a renewable resource forestry in Europe susceptibility of plantation monocultures to insect outbreaks characteristics of insect outbreaks. Part 2 Forests as a habitat for insects: the community of insects on trees climate and weather in the ecology of forest insects biotic factors in the ecology of forest insects. Part 3 Trees as a source of food for insects: nutritional requirements of herbivorous insects nutrient content of trees in relation to structure and function food quality and plant defence insect life cycles in relation to the quantity and quality of food host stress and food quality trees and insect population dynamics implications of insect and tree interactions for forest management. Part 4 The nature of forest insect problems: pest problems in relation to the growing forest special plantations and urban pest problems interactions with other insects and disease organisms the impact of insects on forests and forest trees. Part 5 Forest practices: accidental introduction of pests planning of forestry plantations the post-planning stage - silvicultural techniques for pest management forest diversity and pest outbreaks forest practices and integrated pest management. Part 6 Plant resistance: variation in trees and insects defining resistance resistance mechanisms detecting genetically determined resistance in trees breeding for increased pest resistance host resistance and IPM. Part 7 Biological control: the origin and development of biological control in forestry the potential for biological control in forestry the potential for biological control in forestry the characteristics and use of biological control agents the safety of biological control agents. Part 8 Insecticides: historical aspect of insecticide use the scale of control operations with insecticides insecticides and pest biology the impact of insecticides - pests, natural enemies and environment insecticide technology regulation of insecticide use the costs of insecticides and the economics of their use in forestry. Part 9 Behaviour-modifying chemicals: the role of chemicals in insect behaviour the characteristics of pheromones anti-feedants and repellents. Part 10 Integrated pest management: integrated pest management in forestry developing an integrated approach to forest insect control case histories of integrated pest management in forestry.