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Insect Chemical Ecology Research in the United States Department Of Pest Management Science Pest Manag Sci 59:777–787 (online: 2003) DOI: 10.1002/ps.710 Insect chemical ecology research in the United States Department of Agriculture– Agricultural Research Service†‡§ Jeffrey R Aldrich,1∗ Robert J Bartelt,2 Joseph C Dickens,1 Alan L Knight,3 Douglas M Light4 and James H Tumlinson5 1USDA-ARS Chemicals Affecting Insect Behavior Laboratory, Agricultural Research Center-West, Beltsville, MD 20705, USA 2USDA-ARS National Center for Agricultural Utilization Research, 1815 North University Street, Peoria, IL 61604, USA 3USDA-ARS Yakima Agricultural Research Laboratory, Wapato, WA 98951, USA 4USDA-ARS Western Regional Research Center, Albany, CA 94710, USA 5USDA-ARS Center for Medical, Agricultural and Veterinary Entomology, 1700 SW 23rd Drive, Gainesville, FL 32608, USA Abstract: This multi-author paper reviews current work by USDA-ARS scientists in the field of chemical ecology. Work with pheromones, the discovery and development of the codling moth kairomone, studies on insect–plant interactions and chemically mediated tritrophic plant–insect interactions have led to practical methods for control of important insect pests. Published in 2003 for SCI by John Wiley & Sons, Ltd. Keywords: pheromone; kairomone; green-leaf volatiles; tritrophic interactions; Colorado potato beetle; codling moth; Chrysomelidae 1 INTRODUCTION aggregation pheromone for the Colorado potato beetle Discovery and development of environmentally safe (CPB) are described, semiochemical breakthroughs chemicals to manage the behavior of insect pests with the potential to transform CPB management. and their natural enemies is the primary mission of Finally in Section 5, the now substantial evidence is insect chemical ecologists in the Agricultural Research reviewed showing that insect attack induces plants Service (ARS). Here, ARS investigators from four to, in effect, call parasitoids and predators to their different regions of the country summarize a variety defense–a third trophic level of plant protection of approaches to applied chemical ecology. Research beyond that once imagined by scientists. on insect pheromones has long been an important focus of this effort,1 and pheromones continue to be the cornerstone of insect behavioral manipulation, 2 PHEROMONE CHEMISTRY OF as exemplified in Section 2. The extent to which COLEOPTERANS AND HYMENOPTERANS plants are signal sources guiding insect pests to find Pheromones have become important tools for mon- and accept a potential host has become increasingly itoring and controlling agricultural pest populations. apparent in recent years. Section 3 describes the This technology has been developed most for moths discovery of a particular phytochemical that is and certain beetle groups. However, basic pheromone exploited by codling moth females and males to knowledge is still lacking for many economically find host-plants, and how the synthetic kairomone important insect species, both with respect to chem- is becoming part of the arsenal for codling moth istry and with respect to mate-finding biology. An management. In the first part of Section 4, a series ongoing project at The National Center for Agri- of studies are detailed showing that green leaf volatiles cultural Utilization Research (NCAUR) has been to (GLVs) are common, if not ubiquitous, modulators develop such knowledge and to explore the potential of phytophagous insect behavior. Then, discovery of for using the new pheromones in practical insect a GLV-based plant attractant and the male-produced management. ∗ Correspondence to: Jeffrey R Aldrich, USDA-ARS, Chemicals Affecting Insect Behavior Laboratory, Bldg 007, Room 302, BARC-West Beltsville, MD 20705, USA E-mail: [email protected] †One of a collection of papers on various aspects of agrochemicals research contributed by staff of the Agricultural Research Service of the United States Department of Agriculture, and collated and organized by Drs RD Wauchope, NN Ragsdale and SO Duke ‡This article is a US Government work and is in the public domain in the USA §Sections 2, 3, 4, 5 and 6 of this article were contributed by authors RJB, ALK and DML, JCD, JMT and JRA, respectively (Received 25 March 2002; revised version received 18 November 2002; accepted 27 January 2003) Published in 2003 for SCI by John Wiley & Sons, Ltd. 777 JR Aldrich et al Significant progress has been made over the last 2.1 Sap beetles few years on a number of previously unstudied The sap beetle, Coleopterus truncatus Randall pheromone systems. A typical approach has involved (Coleoptera: Nitidulidae), is a known vector of gas chromatographic (GC) comparison of volatiles Ceratocystis fagacearum (Bretz) Hunt, the fungus that collected from males and females combined with causes the oak wilt disease. The beetles are minute electroantennographic detector (GC-EAD) analysis of in size and highly cryptic in habits, and a pheromone these extracts. Sex-specific compounds that stimulate for this species would be useful for monitoring its intense antennal responses are considered likely abundance and field activity patterns, particularly with pheromone components. These compounds are respect to disease transmission. identified by spectrometric methods (primarily MS Collections of volatiles from groups of males and and NMR) and are then synthesized. The synthetic females feeding on a wheat-germ/brewer’s-yeast arti- compounds are subsequently used to verify behavioral ficial diet revealed three male-specific compounds activity, typically in the field. The increased sensitivity that produced strong responses from the antennae of modern analytical methods has made it possible of both sexes. These were identified as methyl- to detect, evaluate antennal activity and identify key branched, unsaturated hydrocarbons (1–3, Fig 1), compounds even if only small numbers of individuals and the synthesized compounds were attractive to are available (in favorable cases, fewer than one both sexes in field tests, supporting their characteriza- hundred). Sufficient numbers of insects can often tion as an aggregation pheromone.2 Nitidulids of the be obtained from field collections, thus obviating genus Carpophilus also use alkyl-branched unsaturated labor-intensive rearing programs. Recent results for hydrocarbons as aggregation pheromones.3 An impor- five pheromone systems are summarized below. tant feature for both genera is that the pheromones Coleoptera: Nitidulidae Colopterus truncatus (males) 123 Coleoptera: Chrysomelidae Phyllotreta cruciferae (males) (4-9) Aphthona flava, A czwalinae, A cyparissiae (males) (4-11) H H H H H H H H 4 5 6 7 H H H H H H H HO O HO 8910 11 Coleoptera: Chrysomelidae Hymenoptera: Cephidae Oulema melanopus (males) Janus integer (females) H OH O O (CH3)4 (CH2)7CH3 O 12 13 Hymenoptera: Cephidae Cephus cinctus (both sexes, but primarily males) O OO (CH2)8 H (CH2)8 (CH2)n O O O 15: n = 12 O 14 16: n = 14 17: n = 16 CH3(CH2)7 H OH O 18 O 19 Figure 1. Structures of insect-derived compounds recently identified at The National Center for Agricultural Utilization Research (NCAUR). The producing sex is noted in parentheses. 778 Pest Manag Sci 59:777–787 (online: 2003) Insect chemical ecology act synergistically with the volatiles emitted by various and enough material was accumulated for spectral host-related yeasts and fungi. analysis (NMR was essential). The compound was identified as a hydroxyketone (12, Fig 1) and was 2.2 Flea beetles synthesized.6 Future research will include field testing Flea beetles (Coleoptera: Chrysomelidae, subfamily of this compound. Alticinae) are a large group that includes both serious crop pests and beneficial species. Adults of most 2.4 Currant stem girdler sawfly species feed on host plant foliage, while the larvae The currant stem girdler (Janus integer Norton, bore into the roots. No pheromone structures had Hymenoptera: Cephidae) is an occasional pest 4 been published for this group, but Peng et al reported of red currant (Ribes spp) in North America that males of the crucifer flea beetle, Phyllotreta and is of economic importance in the state of cruciferae (Goeze), feeding on host foliage (Brassica Washington. Pheromones in the Cephidae were napus L) attracted both males and females in the previously unexplored. Collections of volatiles from field, indicating the existence of a male-produced males and females were analyzed by GC-EAD. A aggregation pheromone. This species is a major female-specific compound was detected that caused an pest of rapeseed and canola in the prairies of the intense response in male (but not female) antennae, a northern USA and Canada. Chemical studies were pattern indicative of a sex pheromone. The compound begun at NCAUR with P cruciferae and, subsequently, was identified as a lactone (13, Fig 1), and the racemic with three beneficial species of Aphthona [A flava form was synthesized.7 This synthetic compound was Guillebeau, A czwalinae (Weise), and A cyparissiae found to be attractive to males in the initial field test (Koch)], which had been introduced into the USA (James DG, pers comm). The natural compound was as biological control agents of leafy spurge (Euphorbia subsequently determined to be the (R) enantiomer esula L), a serious rangeland weed. (shown in Fig 1) by chiral GC and chiral synthesis Comparisons of collections of volatiles from males (Petroski RJ, pers comm). and females of P cruciferae feeding on canola or cabbage revealed six male-specific compounds, which 2.5 Wheat stem sawfly were identified as the sesquiterpenoids 4–9 in Fig 1.5 The wheat stem sawfly, Cephus cinctus Norton Intriguingly, males of all three of the Aphthona (Hymenoptera: Cephidae), is a serious pest of winter species also emitted compounds 4–9,plustwo wheat (Triticum aestivum L) in the northern Great other compounds (10 and 11, Fig 1) not detected Plains of the USA and Canada. GC-EAD analysis from P cruciferae. All except 7 and 8 are new of collections of volatiles from males and females natural products. Ratios of the compounds were revealed a situation very different from that with the quite consistent within species but differed between cephid, Jinteger (Section 2.4).
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