Preference of Neoseiulus Californicus (Acari: Phytoseiidae) for Volatiles
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Revista Brasileira de Entomologia 63 (2019) 283–289 REVISTA BRASILEIRA DE Entomologia A Journal on Insect Diversity and Evolution www.rbentomologia.com Biological Control and Crop Protection Preference of Neoseiulus californicus (Acari: Phytoseiidae) for volatiles of Bt maize induced by multiple herbivory Jéssica Mayara Coffler Botti , Mayara Loss Franzin , ∗ Marcos Antonio Matiello Fadini , Júlio Onésio Ferreira Melo Universidade Federal de São João Del Rei, Campus Sete Lagoas, Sete Lagoas, MG, Brazil a b s t r a c t a r t i c l e i n f o Article history: Plant indirect induced defenses against herbivores are characterized by the production of plant volatiles Received 9 April 2019 that to attract natural enemies. The objective of this study was to evaluate whether the attack of the Accepted 23 September 2019 two-spotted spider mite Tetranychus urticae or the multiple herbivory of T. urticae together with the Available online 17 October 2019 fall armyworm Spodoptera frugiperda are able to elicit indirect induced defense in conventional and Bt Associate Editor: Mariane A. Nickele maize plants. The experiment was carried out in the laboratory using Y-tube olfactometer, evaluating the predatory mite Neoseiulus californicus olfactory preference for plant volatiles. The treatments involved: Keywords: Clean Conventional Plant; Conventional Plant Infested with T. urticae; Clean Bt Plant; Bt Plant Infested Indirect induced defense with T. urticae; Conventional Plant Infested with T. urticae + fall armyworm; Bt Plant Infested with T. Tetranychus urticae urticae + fall armyworm. For the chemical analyzes the Trace GC Ultra gas chromatograph was used, Secondary plant compounds paired with the Polaris Q mass spectrometer, GC MS system. Neoseiulus californicus was also unable to Plant resistance Fall armyworm distinguish between volatiles from both conventional and Bt infested maize plants. Moreover, there was no discrimination of the predator mite between plants under single and multiple infestations, both in conventional and Bt maize. When comparing conventional and Bt plants, both with multiple infestation, the predator mite N. californicus had no preference among these sources of odors. However, there was observed chemical changes of the volatiles among the groups of plants studied. Thus, it is suggested that the groups of plants under study have chemical modifications, but they are not able to attract N. californicus. In addition, Bt plants response was similar to conventional plants on attracting N. californicus. © 2019 Published by Elsevier Editora Ltda. on behalf of Sociedade Brasileira de Entomologia. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/ ). Introduction consequently exposed (Torres and Ruberson, 2008). Although the use of genetically modified plants has advantages (Kouser and The plants developed a diversity of defense mechanisms against Qaim, 2011), the impact of these plants on the foraging behavior different conditions, such as herbivorous attack (Bergelson et al., of natural enemies has been questioned (Poppy and Sutherland, 2001). A mechanism against herbivores is the indirect induced 2004). Hence, it is important to understand the influence of Bt plant defense, characterized by the production of plant volatiles after proteins on non-target organisms. herbivory that are able to attract natural enemies of herbivores Further, the cultivation of Bt plants promotes a decrease in the (Arimura et al., 2005; Schaller, 2008). use of insecticides (Cattaneo et al., 2006). In this way, species of non- In addition to induced defenses, plants have constitutive target pests of lesser economic importance, such as Tetranychus defenses that are expressed continuously and independent of her- urticae (Koch) (Acari: Tetranychidae) in maize, may become more bivorous attack (Price et al., 1980). One example is Bt plants, since important, because the lower use of insecticides applied to target they continuously express the defense protein. For example, 30F35 pests may increase their population. Hx maize expresses the Cry1F protein that confers lepidopteran Therefore, it is important to understand if Bt-based corn plants resistance. However, not only the target herbivores are exposed are capable able of releasing volatiles to attract the natural ene- to toxins, non-target herbivores and their natural enemies are mies of T. urticae. This questioning is performed because indirect ∗ Corresponding author at: Universidade Federal de São João Del Rei. MG 424, km 47. Sete Lagoas, MG, 35701-970, Brazil. E-mails: [email protected] (J.M. Botti), [email protected] (M.L. Franzin), [email protected] (M.A. Fadini), [email protected] (J.O. Melo). https://doi.org/10.1016/j.rbe.2019.09.003 0085-5626/© 2019 Published by Elsevier Editora Ltda. on behalf of Sociedade Brasileira de Entomologia. This is an open access article under the CC BY-NC-ND license (http:// creativecommons.org/licenses/by-nc-nd/4.0/). 284 J.M. Botti et al. / Revista Brasileira de Entomologia 63 (2019) 283–289 induced defense has costs for the plants (Steppuhn and Baldwin, The group of plants infested with T. urticae + fall armyworm, 2008). To trigger this defense, the plant needs to allocate physiolog- besides the infestation of the mite that was identical to the previous ical and ecological resources, which can affect relevant processes of group, a third instar larvae was isolated with a “clip cage”. This was the plant, such as growth and reproduction (Steppuhn and Baldwin, carried out on a different leaf of T. urticae infestation, also 20 days 2008). after sowing. A second concerns is the interaction among different types of plant defenses mechanisms (Kant et al., 2015; Price et al., 1980). Olfactometer test Studies have demonstrated significant differences in the amount of volatiles emitted between Bt and non-Bt maize plants (Dean and Plants after 48 h of infestation were used to carry out the tests in De Moraes, 2006; Turlings et al., 2005). This shows that plants with Y-tube olfactometer, with the following treatments: Clean Conven- constitutive defenses (e.g. Bt) can alter the production of volatiles tional Plant X Conventional Plant Infested with T. urticae; Clean Bt and, consequently, the attraction of natural enemies. This suggests Plant X Bt Plant Infested with T. urticae; Conventional Plant Infested a negative interaction between these defenses (Shimoda, 2010). with T. urticae X Bt Plant Infested with T. urticae; Conventional Plant Furthermore, another factor that can alter induced defense is Infested with T. urticae X Conventional Plant Infested with T. urticae the multiple attacks of herbivore species, affecting preference and + fall armyworm; Plant Bt Infested with T. urticae X Bt Plant Infested performance of herbivores and of their natural enemies (Dicke with T. urticae + fall armyworm; Conventional Plant Infested with et al., 2009). In nature, plants are not exposed to a single herbi- T. urticae + fall armyworm X Plant Bt Infested with T. urticae + fall vore. The attack of multiple herbivores can alter the emission of armyworm. Each treatment consisted of three true replicates (i.e. compounds, both positively and negatively in the attractiveness of odor sources) then 15 responses of predatory mites were obtained natural enemies (Kaplan et al., 2009; Rodriguez-Saona et al., 2005; from each replicate, totaling 45 releases of predatory mites per Viswanathan et al., 2005). This work evaluated whether both con- treatment. ventional and Bt maize plants induce the production of volatile A Y-tube olfactometer was used to study the response of N. cal- compounds attractive to Neoseiulus californicus (McGregor) (Acari: ifornicus. The glass Y-Tube (with dimensions of 21.0 cm of length Phytoseiidae) after herbivory of T. urticae. And if conventional and of each arm and 3.5 cm of diameter) had a Y-shaped metal wire Bt maize plants induced by simple (T. urticae) and multiple her- inside to facilitate the movement of mites. Containers consisting bivory (T. urticae + fall armyworm), produce volatile compounds of a plastic box were attached to the Y-tube both arms. The Y- attractive to the predatory mite N. californicus. tube base was connected to a vacuum pump that directed the flow from the containers to the Y-tube arms. A flowmeter was connected Materials and methods to the containers air outlet and was used to calibrate the air inlet speed in each olfactometer arm. The wind speed was 0.50 m/s on Rearing of two-spotted spider mite, fall armyworm and predatory each Y-tube arm. The predators were individually introduced at the mite starting point on the steel wire and then observed until a predator reached 1/3 of an arm, where the mite response was considered. Infested leaves of Sorghum plants with T. urticae were col- Predatory mites who did not pass 1/3 of either arm within 5 min lected in the greenhouse at Embrapa Milho e Sorgo facilities, and were accounted unresponsive¨ ¨. After every five responses, the odor taken to the Entomology Laboratories of the Universidade Fed- sources positions were reversed. eral de São João Del-Rei (UFSJ), in Sete Lagoas, Minas Gerais, Brazil. To establish the colony, adult mites were collected, individ- Chemical analysis of plant volatiles ually using a stereomicroscope and a hairs brush, then transferred to jack bean plants (Canavalia ensiformis) with two completely For chemical analysis, a leaf sample of 0.250 g of each plant expanded cotyledon leaves and without any phytosanitary treat- was taken. The chemical analysis was performed using solid phase ment. microextraction (SPME) in headspace mode and Trace GC Ultra The bean plants used were planted in 1.0 L plastic pots using gas chromatograph (ThermoScientific, San Jose, CA) paired with ® Terral Solo substrate. Infested plants were kept isolated in the Polaris Q mass spectrometer (ThermoScientific, San Jose, CA), screened cages in order to prevent external infestation with other GC–MS system, with an ion-trap type analyzer (Garcia et al., 2016).