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319 SUSTANCIAS VOLÁTILES DE GRANOS DE ARROZ INDUCIDAS POR HERVÍBOROS AFECTAN LA ATRACCIÓN DE ENEMIGOS HERBÍVOROS Qingfeng Tang, Tiansheng Yang Y Junqi Jiang RESUMEN

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319 SUSTANCIAS VOLÁTILES DE GRANOS DE ARROZ INDUCIDAS POR HERVÍBOROS AFECTAN LA ATRACCIÓN DE ENEMIGOS HERBÍVOROS Qingfeng Tang, Tiansheng Yang Y Junqi Jiang RESUMEN HERBIVORE-INDUCED RICE GRAIN VOLATILES AFFECT ATTRACTION BEHAVIOR OF HERBIVORE ENEMIES QingFeng Tang, TianSheng Yang and JunQi Jiang SUMMARY The responses of female Lariophagus distinguendus (Hy- vae damaged grains. Further experiments revealed the expe- menoptera: Pteromalidae) to volatile signals derived from rienced L. distinguendus females were apparently attracted herbivore-induced rice grain was investigated in the static by the odors released from rice induced by the saliva of the four-chamber olfactometer. Results showed that the L. dis- weevil larvae. Moreover, we compared the responses of fe- tinguendus females, irrespective of experience, were appar- male parasitoids to odors released from rice induced by pro- ently attracted by the odors released from rice grain. The tein substances of regurgitant and non-protein substances. active chemicals in the saliva of the maize weevil, Sitophi- The experiments revealed that volatiles attracting parasitoid lus zeamais, that elicit the production of rice volatiles was L. distinguendus are possibly induced by the some protein investigated. Results showed that artificially damaged grains substances from weevil larvae saliva. The effect of experi- do not emit large amounts of volatiles that attract the ex- ence on the response of the parasitoids to several stimuli perienced parasitoid females emitted by the S. zeamais lar- was discussed. Introduction researchers to evaluate other option is effective in con- particularly for foliage. It is alternative reduced-risk con- trolling insect pests of stored well documented that carniv- In many parts of the world trol methods for stored-grain products and has a positive orous arthropods often use rice is the major component protection (Tang et al., 2009). impact on the quality of volatile chemical cues that are of the diet. Unfortunately, a To resolve the above prob- stored cereal products. These released from the food plants large proportion of the annu- lems, new environmental chemicals that are beneficial of their victims when search- al yield is lost during stor- friendly methods to control for both the carnivores and ing for prey or hosts. By way age, and these losses have the insect damages have be- the plant have been termed of comparison, little is known been partly attributed to in- come of interest to research- ‘herbivore induced syno- about the release of HIS from sect pests resulting in the ers; biological control of mones’ (HIS). Since the first reproductive tissue such as deterioration of the quality pests using parasitoids or description of this interaction rice grain (Tang et al., 2009). and the quantity of stored predators is an example between plants, herbivores The present paper is devoted rice grains (Machado et al., (Scholler et al., 1997). The and carnivores mediated by to a tritrophic system consist- 2008). To protect grains from use of natural enemies such HIS there have been numer- ing of the rice grains (Oryza. the insect damages, tons of as parasitoids for the control ous studies on this phenome- sativa L.), larvae of the maize insecticides have been ap- of stored-product pests has non. There is evidence to sug- weevil Sitophilus zeamais plied to control the pests, been suggested for many gest that HIS can regulate tr- (Coleoptera: Curculionidae) that which have resulted in envi- years (Hase, 1924; Ryabov, itrophic interactions among are endophytic in rice grains, ronmental damages, pest re- 1926). However, little is plants, insect herbivores, and and the pteromalid wasp surgence, pest resistance to known about tritrophic inter- natural enemies of the herbi- Lariophagus distinguendus insecticides, and lethal ef- actions among rice grain, vores (Steinberg et al., 1993; (Hymenoptera: Pteromalidae), fects on non-target organ- insect herbivores, and natural Ockroy et al., 2001; Tinzaara an idiobiont ectoparasitoid of isms. Furthermore, because enemies of the herbivores. et al., 2005; Dalessandro beetle larvae, infesting seeds of cost, these pesticides are The use of specific attrac- et al., 2009; Deshpande and and grains. The maize weevil becoming increasingly inac- tants might represent an op- Kainoh, 2012; Peñaflor and S. zeamais is one of the most cessible to farmers, particu- tion to increase the presence Bento, 2013; Desurmont common and destructive stor- larly in developing countries. of the parasitoid wasp in gra- et al., 2015; Yoneya and Miki, age pests of stored cereal This fact, combined with nary or could be coupled with 2015). To date, HIS have been grains in the world. The adult the consumer’s demand for biocontrol programmes to im- demonstrated mostly in the beetles can attack whole residue-free food, prompted prove biocontrol services. The somatic tissue of plants, grains, and eggs are laid KEYWORDS: / Grain Volatiles / Lariophagus distinguendus / Multitrophic Interaction / Olfactometer Tests / Sitophilus zeamais / Received: 11/26/2015. Modified: 03/31/2016. Accepted: 04/04/2016. QingFeng Tang. Ph.D., Anhui TianSheng Yang. B.S., AAU, JunQi Jiang. Ph.D., AAU, Protection, AAU. 130# West Agricultural University (AAU), China. Master Degree Candi- China. Professor, AAU, China. Changjiang Road, Hefei China. Professor, AAU, China. date, AAU, China. Address: Department of En- 230036, P. R. China. e-mail: tomology, School of Plant [email protected] MAY 2016, VOL. 41 Nº 5 0378-1844/14/07/468-08 $ 3.00/0 319 SUSTANCIAS VOLÁTILES DE GRANOS DE ARROZ INDUCIDAS POR HERVÍBOROS AFECTAN LA ATRACCIÓN DE ENEMIGOS HERBÍVOROS QingFeng Tang, TianSheng Yang y JunQi Jiang RESUMEN En un olfatómetro de cuatro cámaras se estudiaron las res- por las larvas de S. zeamais de granos dañados. Experimen- puestas de hembras de Lariophagus distinguendus (Hymenop- tos adicionales revelaron que las hembras experimentadas de tera: Pteromalidae) a señales volátiles provenientes de granos L. distinguendus eran aparentemente atraídas por los olores de arroz e inducidas por herbívoros. Los resultados mostraron desprendidos de los granos de arroz e inducidos por la saliva que las hembras de L. distinguendus, independientemente de de las larvas de gorgojos. Adicionalmente se compararon las su experiencia, son atraídas por los olores despedidos por los respuestas de los parasitoides hembra a olores emanados del granos de arroz. Se investigaron las sustancias activas presen- arroz inducidas por proteínas del vómito y por sustancias no tes en la saliva del gorgojo del maíz, Sitophilus zeamais, que proteicas. Los experimentos mostraron que los productos vo- inducen la producción de sustancias volátiles en el arroz. Los látiles que atraen los parasitoides de L. distinguendus son in- resultados mostraron que los granos de arroz artificialmente ducidos probablemente por algunas sustancias proteicas de la dañados no emiten cantidades de sustancias volátiles capa- saliva del gorgojo. Se discute el efecto de la experiencia previa ces de atraer a parasitoides hembras experimentadas emitidas en la respuesta de los parasitoides a diversos estímulos. SUBSTÂNCIAS VOLÁTEIS DE GRAÕS DE ARROZ INDUZIDAS POR HERVÍBOROS AFETAM A ATRAÇÃO DE INIMIGOS HERBÍVOROS QingFeng Tang, TianSheng Yang e JunQi Jiang RESUMO Em um olfatômetro de quatro câmaras se estudaram as res- zeamais de grãos danificados. Experimentos adicionais revela- postas de fêmeas de Lariophagus distinguendus (Hymenoptera: ram que as fêmeas experimentadas de L. distinguendus eram Pteromalidae) a sinais voláteis provenientes de grãos de arroz aparentemente atraídas pelos odores desprendidos dos grãos e induzidas por herbívoros. Os resultados mostraram que as de arroz e induzidos pela saliva das larvas de gorgulhos. Adi- fêmeas de L. distinguendus, independentemente de sua experi- cionalmente se compararam as respostas dos parasitoides fê- ência, são atraídas pelos odores desprendidos pelos grãos de meas a odores emanados do arroz induzidas por proteínas do arroz. Investigaram-se as substancias ativas, presentes na sa- vômito e por substancias não proteicas. Os experimentos mos- liva do gorgulho do milho, Sitophilus zeamais, que induzem à traram que os produtos voláteis que atraem os parasitoides produção de substâncias voláteis no arroz. Os resultados mos- de L. distinguendus são induzidos provavelmente por algumas traram que os grãos de arroz artificialmente danificados não substâncias proteicas da saliva do gorgulho. Discute-se o efei- emitem quantidades de substâncias voláteis capazes de atrair to da experiência prévia na resposta dos parasitoides a diver- parasitoides fêmeas experimentadas emitidas pelas larvas de S. sos estímulos. inside kernels in which the volatiles on the taxis behavior relative humidity (r.h.) and a Insects for bioassays larvae feed and develop entire- of L. distinguendus and its photoperiod of L14: D10. To ly and pupate. The oviposition mechanisms. Such research was rear L. distinguendus, 50 newly Parasitoids used in experi- hole is closed with an egg plug of importance to reveal the emerged adult wasps were ments were about 3 days old. (Hill, 1983; Storey, 1987; mechanism of indirect defense placed into Petri dishes (9cm To obtain naive parasitoids for Kouninki et al., 2007). L. dis- of rice grains against larvae of diameter, 1cm high) with ~50g bioassays, freshly emerged male tinguendus has been suggested S. zeamais and, it was also very of rice grains infested by 3rd- and female parasitoids were as a biological control agent important in the understanding 4th instar larvae of S. zeamais collected from the infested against the maize weevil in of the ecological functions
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