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Present Or Past Herbivory: a Screening of Volatiles Released from Brassica Rapa Under Caterpillar Attacks As Attractants for the Solitary Parasitoid, Cotesia Vestalis

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Present Or Past Herbivory: a Screening of Volatiles Released from Brassica Rapa Under Caterpillar Attacks As Attractants for the Solitary Parasitoid, Cotesia Vestalis J Chem Ecol (2010) 36:620–628 DOI 10.1007/s10886-010-9802-6 Present or Past Herbivory: A Screening of Volatiles Released from Brassica rapa Under Caterpillar Attacks as Attractants for the Solitary Parasitoid, Cotesia vestalis Soichi Kugimiya & Takeshi Shimoda & Jun Tabata & Junji Takabayashi Received: 20 February 2010 /Revised: 7 April 2010 /Accepted: 11 May 2010 /Published online: 20 May 2010 # Springer Science+Business Media, LLC 2010 Abstract Females of the solitary endoparasitoid Cotesia decreased after removal of the host larvae, whereas vestalis respond to a blend of volatile organic compounds terpenoids and their related compounds continued to be (VOCs) released from plants infested with larvae of their released at high levels. Benzyl cyanide and dimethyl host, the diamondback moth (Plutella xylostella), which is trisulfide attracted parasitoids in a dose-dependent manner, an important pest insect of cruciferous plants. We investi- whereas the other compounds were not attractive. These gated the flight response of female parasitoids to the results suggest that nitrile and sulfide compounds tempo- cruciferous plant Brassica rapa, using two-choice tests rarily released from plants under attack by host larvae are under laboratory conditions. The parasitoids were more potentially more effective attractants for this parasitoid than attracted to plants that had been infested for at least 6 hr by other VOCs that are continuously released by host- the host larvae compared to intact plants, but they did not damaged plants. distinguish between plants infested for only 3 hr and intact plants. Although parasitoids preferred plants 1 and 2 days Key Words Herbivore-induced plant volatiles . after herbivory (formerly infested plants) over intact plants Indirect defense . Tritrophic interaction . Brassica rapa . they also preferred plants that had been infested for 24 hr Plutella xylostella . Cotesia vestalis over formerly infested plants. This suggests that parasitoids can distinguish between the VOC profiles of currently and formerly infested plants. We screened for differences in Introduction VOC emissions among the treatments and found that levels of benzyl cyanide and dimethyl trisulfide significantly Many plant species release specific blends of volatile organic compounds (VOCs) in response to attack by herbivores. The release of VOCs induced by herbivory can attract natural enemies of herbivores and may guide : parasitoids or predators to their hosts or prey (Turlings et al. S. Kugimiya (*) J. Tabata 1990; Takabayashi and Dicke 1996). Chemical information National Institute for Agro-Environmental Sciences, of this nature has attracted considerable attention as an Kannondai 3-1-3, Tsukuba, Ibaraki 305-8604, Japan indirect defense of plants against herbivorous insects e-mail: [email protected] mediated by their natural enemies at a higher level in a tritrophic interaction system (Karban and Baldwin 1997; T. Shimoda Sabelis et al. 2007). However, the plant defense may not be National Agricultural Research Center, Kannondai 3-1-1, constant, since the composition of plant VOCs can vary Tsukuba, Ibaraki 305-8666, Japan both in quality and in quantity, depending on various biotic and abiotic factors, and these changes can affect the J. Takabayashi attractiveness of the plants to natural enemies of the Center for Ecological Research, Kyoto University, Hirano 2-509-3, herbivores (Takabayashi et al. 1994; Maeda et al. 2000; Otsu, Shiga 520-2113, Japan Gouinguené and Turlings 2002). J Chem Ecol (2010) 36:620–628 621 The profile of VOCs also can change during the course after the start of herbivory and after removal of the larvae, of herbivory (Loughrin et al. 1994; Turlings et al. 1998; and tested synthetic versions of VOCs to evaluate their Scascighini et al. 2005). Diurnal cycles in the release of effectiveness as attractants for parasitoids. VOCs are related to the attraction of natural enemies of herbivores or to deterrence of herbivores (Loughrin et al. 1994; De Moraes et al. 2001; Shiojiri et al. 2006a), and the Methods and Materials importance of the effects of light in a diurnal cycle have been reported (Gouinguené and Turlings 2002). However, Plants and Insects Japanese mustard spinach, Brassica there is a little information on changes in VOCs of infested rapa L. var. perviridis (Capparales: Brassicaceae), was plants during and after herbivory, particularly from the cultivated in a greenhouse (25±3°C, 60±10% relative perspective of potential differences in plant-insect inter- humidity (RH), 16L:8D photoperiod). Five plants were actions between the two phases (Mattiacci et al. 2001; reared in a plastic pot (90 mm diam, 70 mm depth) for 4– Hoballah and Turlings 2005). In natural ecosystems, 5 wk, and were used for both insect rearing and flight herbivores may not stay on a single plant, but may move preference tests. Diamondback moths, P. xylostella, origi- among plants, be eliminated by other predators (Shiojiri and nally collected from fields in Ayabe, Kyoto Prefecture, Takabayashi 2005), or pupate and become unsuitable Japan, in 2001, were mass-reared on potted plants in a targets for parasitoids to oviposit. Thus, parasitoids may climate-controlled room (25±3°C, 60±10% RH, 16L:8D be able to distinguish between plants currently being photoperiod). Eggs were collected every day, and hatched attacked (hereafter, “infested plants”) and plants that were larvae were reared on cut plants in small cages (width formerly attacked by herbivores (hereafter, “formerly 25 cm, depth 15 cm, height 10 cm). The solitary parasitoids infested plants”). By focusing on the difference in compo- C. vestalis, which parasitize mainly P. xylostella larvae, sitions of VOCs released from infested and formerly were obtained from their hosts collected in the same fields infested plants, we may be able to identify effective and were reared on P. xylostella-infested plants under the attractants for parasitoids. same conditions as their hosts. For use in the two-choice In Japan, Brassica plants (Capparales: Brassicaceae) are test, wasp cocoons were collected from the stock culture, grown during the spring, close enough together for adjacent and newly emerged adults were maintained with 50% plants to touch. A tritrophic system forms among the plant aqueous honey in acryl cages (width 35 cm, depth 25 cm, species, herbivorous insects, and their parasitoids. Cater- height 30 cm) separately from the host-infested plants until pillars of the diamondback moth, Plutella xylostella L. the experiments. (Lepidoptera: Yponomeutidae), are oligophagous on crucif- erous plants. The specialist parasitoid wasp Cotesia vestalis Flight Response of Parasitoids to Host-infested Plants and (Haliday) [= C. plutellae (Kurdjumov)] (Hymenoptera: to Synthetic Compounds The flight response of C. vestalis Braconidae) oviposits on P. xylostella larvae. The female females was assessed by using a two-choice test in an acrylic parasitoids are attracted to crucifers infested by P. xylostella chamber (width 35 cm, depth 25 cm, height 30 cm, 3,000 larvae through a specific blend of VOCs (Shiojiri et al. lux) in a climate-controlled room (25±3°C, 60±10% RH) 2000) and lay a single egg per host. (Shiojiri et al. 2000). Infested plants were prepared by Under laboratory conditions, infested cruciferous plants allowing 15 third-instar P. xylostella to feed on potted plants attract various parasitoids by emitting specific blends of of B. rapa for 3, 6, and 24 hr (hereafter, 3, 6, and 24 hr- VOCs (Mattiacci et al. 1994; Shiojiri et al. 2000;Van infested plants, respectively; Fig. 1). The infestation began at Poecke et al. 2001), and the mechanisms for the induction, 12:00 am, during the middle of the light phase of the regulation, and biosynthesis of those VOCs have been photoperiod. Formerly infested plants were prepared by characterized (Mattiacci et al. 1995; Van Poecke and Dicke removing the host larvae from 24 hr-infested plants and then 2002; Shiojiri et al. 2006b;D’Auria et al. 2007; Herde et al. by using the plants for experiments 48 and 72 hr after the 2008). However, it is unclear which VOC components start of the infestation (1 and 2d-after plants, respectively; attract C. vestalis. In the present study, we screened for Fig. 1). In every treatment, we used almost the same size potential parasitoid attractants, by investigating the flight larvae, showing similar feeding activity, and the areas of response of female parasitoids to plants that had been damage caused by the larvae were identical for a given infested by their host larvae, P. xylostella, for different 24 hr-infested plant and the 1d-after and 2d-after versions of durations (infested plants), and their response to plants at that plant. Intact plants were prepared as controls, without various times after herbivory (formerly infested plants). We any treatment. To produce a two-choice test design, one then analyzed the headspace volatiles released from potted plant from a given treatment was placed ca. 10 cm infested and formerly infested plants to establish the time from a plant that had been subjected to a different treatment course of changes in the relative amounts of plant VOCs inside the acrylic chamber, with the leaves not overlapping. 622 J Chem Ecol (2010) 36:620–628 Hosts attacking Collection of Volatiles Headspace volatiles were sampled Hosts removed from whole potted plants subjected to different treatments 2d-after plants (infested plants, formerly infested plants, and intact plants) 1d-after plants using a dynamic headspace collection system in a climate- 24h-infested plants controlled room (25±3°C, 60±10% RH, 16L:8D photope- 6h-infested plants riod). Each pot of plants was placed in a 2-l glass container, and the container was sealed with a glass lid that contained 3h-infested plants an air inlet and an air outlet. The container then was tightly 0h-infested plants (Intact plants) sealed with metal clamps on the lid. Incoming air was 7248 246 3 0 purified by filtration through silica gel and activated charcoal, and was actively pumped through the container Time (h) at a flow rate of 300 ml min−1, constantly monitored and Fig.
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