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- ArticleMeade HM, Signer ER.Proc Natl Acad Sci U S A. 1977 May;74(5):2076-8.The drug resistance factor RP4, originally isolated in Pseudomonas, was transferred to Rhizobium meliloti. In that strain, RP4 promotes conjugational transfer of chromosomal markers to form haploid recombinants. This mating system has been used to construct a linkage map of R. meliloti.