The perception of IGP risk by T. rapae from M. brunneum

but not from B. bassiana may relate to differences in cues emitted by the two fungi. However, these cues may be dependent on the context and complexity of the tested system which may not have been reflected by our experimental arenas. Additional studies should expand on the complexity of our system in order to provide a more complete volatile exposure. For vegetable cruciferous crops, mixing entomopathogenic fungi into the substrate when raising plantlets in the greenhouse for subsequent transplanting to the field would be a convenient method for control of the inoculum levels applied. Chandler and Davidson, 2005 and Razinger et al., 2014 found that this method provided some control of D. radicum using Metarhizium

sp. Seed treatment may be another approach since Keyser ZD1839 manufacturer et al. (2014) found that seed treatment by M. brunneum (isolate EPZ015666 KVL 04-57 as in this study) resulted in infection in insects exposed to the growing roots. These two methods would also take advantage of the endophytic and rhizosphere competent property of Metarhizium sp. ( Sasan and Bidochka, 2012, Razinger et al., 2014 and Wyrebek et al., 2011) in order for the fungi to preestablish before D. radicum attack. This study demonstrated that the tested M. brunneum isolate is a promising biological control candidate against D. radicum larvae. Furthermore, T. rapae showed an ability to perceive and react to the IGP risk posed by M. brunneum while B. bassiana was not avoided to the same extent. Thus M. brunneum has the potential to be used for biological control against D. radicum with a low expected risk to T. rapae populations. The potentially complementary biological control effect against immature D. radicum by conservation biological control targeting T. about rapae populations in combination with inoculation with M. brunneum must be studied under field conditions. We are grateful for the advice and technical

assistance from Dr. Lorna Migiro, technicians Louise Lee Munk Larsen and Mira Rur, entomologist Britt Åhman and the student Laura Engel. We are indebted to Dr. Mario Porcel for statistical discussions, Dr. Ulf Nilsson and Chad Alton Keyser for valuable manuscript comments, and furthermore C.A.K. for language editing. We would like to thank Sebastien Dugravot, University of Rennes 1, for providing the initial strain of T. rapae and Rosemary Collier, University of Warwick, for providing the start culture of D. radicum. This study was supported by a Ph.D. grant to L.-M.R. through the financers Swedish Research Council for Environment, Agricultural Sciences and Spatial Planning (FORMAS; project number 2009-5824-14994-47) and the Swedish University of Agricultural Sciences (SLU), for the SLU affiliated scientists, and by University of Copenhagen for N.V.M.