Linkage group B06 also has few major R-genes [9], with the notable exception of Ur-4, despite its apparent abundance of RGH sequences. The position of bc-3 was not considered, as this is a recessive R-gene that has been suggested to be related to a family of elongation initiation factors [56]. However, the Ur-4 gene, as well as a QTL, for white mold [9] was observed to lie in the same region as BMr51 to BMr302. Linkage group B07 contained Phs, a phaseolin-encoding

locus associated with a common bacterial blight QTL, as well as 4 RGH-SSR plus RGH4b, in the region suspected to contain the R-genes (Co-5, Co-6) and further QTL for anthracnose and common bacterial blight resistance. The three

R-genes and multiple QTL on linkage group B08 aligned well with RGH genes. Co-4, although suspected to be a protein kinase gene, was near the loci RGH15a and RGH15c along with QTL for common Cyclopamine manufacturer Fulvestrant cost bacterial blight and white mold resistance. QTL for resistance to the same diseases plus a QTL for anthracnose resistance were near RGH2, BMr244, and BMr269 and a previously unmapped RGH-RFLP named EcoRV334, which was in the region containing the Phg-2 (angular leaf spot) and Ur-13 (rust) resistance genes [9]. The next two linkage groups were contrasting, in that B09 had few RGH-SSR (3) and few QTL for resistance, while B10 had a large number of RGH genes (10) and many QTL for various diseases. Linkage group B10 has emerged as being very important for angular leaf spot resistance. One report cites anthracnose resistance in the middle of B10 although this is unconfirmed

in other studies [34]. Major R-genes for angular leaf spot on B10 could be analyzed Cyclin-dependent kinase 3 in relation to Phg-1, a new Andean R-gene on B01 [57]. The final chromosome-linkage group B11, especially the end of the long arm, has been long known to be a hotspot for R-genes [9]. From the bottom of B11, there was alignment of BMr207 and RGH1a with Co-2, Ur-3, Ur-11, and Ur-Dorado [9]. Two other major R-genes for rust, Ur-6 and Ur-7, along with common bacterial blight and web blight QTL, are likely to be tagged by 5 RGH-SSR markers in a more proximal location on the chromosome B11 and in the upper part of the linkage group B11 another QTL for common bacterial blight may align with marker BMr281. In summary, this work established the position of new RGH-SSR markers relative to known R-genes. A large number of RGH-related markers have been developed, including 32 from the BAT93 × Jalo EEP558 population [48], 21 from the Dorado × XAN 176 population [50], and 14 from the Calima × Jamapa population [58]. Mutlu et al. [59] coincidentally mapped 32 RGAP bands in the first of these populations and also detailed alignment with QTL and R-genes.