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Enhanced herbicide metabolism mediated by cytochrome P450s has been proposed as one of the major mechanisms of resistance to fenoxaprop- P -ethyl in a metabolic-herbicide-resistant biotype of Asia minor bluegrass ( Polypogon fugax  Nees ex Steud.). Upon pre-treatment with the P450 inhibitor piperonyl butoxide, a remarkable reduction in metabolic rates of the phytotoxic fenoxaprop- P  has been observed in the resistant plants, implying that constitutive and/or fenoxaprop- P -ethyl-induced up-regulation of specific P450 isoforms are involved in the fenoxaprop- P -ethyl resistance. However, which P450 gene(s) were responsible for the metabolic resistance is still unknown. In this present study, based on the abundant gene resources of  P. fugax  established previously, a total of 48 putative P450 genes were isolated from the metabolic-herbicide-resistant plants and used for gene expression analysis. The most suitable reference genes for accurate normalization of real-time quantitative PCR data were first identified in  P. fugax  and recognized as actin ( ACT ), 18S rRNA ( 18S ), and ribulose-1,5-bisphosphate carboxylase oxygenase ( RUBP ) under fenoxaprop- P -ethyl stress, glyceraldehyde-3-phosphate dehydrogenase ( GAPDH ) and elongation factor 1α ( EF1α ) under mesosulfuron-methyl stress, and  ACT ,  EF1α , eukaryotic initiation factor 4a ( EIF4A ), and 25S rRNA ( 25S ) at different growth stages. Expression analysis of the putative P450 genes revealed that six genes, respectively, annotated as  CYP709B1 ,  CYP71A1-4 ,  CYP711A1 ,  CYP78A9 ,  P450-11 , and  P450-39  were up-regulated more than 10-fold in the resistant plants by fenoxaprop- P -ethyl treatment, and all of them exhibited constitutively and/or herbicide-induced higher transcript levels in the fenoxaprop- P -ethyl-resistant than in the susceptible plants. Three genes, respectively, annotated as  CYPRO4 ,  CYP313A4 , and  CYP51H11  constantly up-regulated in the resistant than in the susceptible plants after fenoxaprop- P -ethyl treatment. Up-regulated expressions of these specific P450 genes were consistent with the higher P450 contents determined in the resistant plants. These results will help to elucidate the mechanisms for P450-mediated metabolic-herbicide resistance in  P. fugax  as well as other grass weed species.

Expression Pattern of Entire Cytochrome P450 Genes and Response of Defenses in a Metabolic-Herbicide-Resistant Biotype of Polypogon fugax