The Evolutionary Moulding in plant-microbial symbiosis: matching population diversity of rhizobialnodA and legumeNFR5genes
AbstractWe propose the Evolutionary Moulding hypothesis that population diversities of partners in nitrogen-fixing rhizobium-legume symbiosis are matched, and tested it in nucleotide polymorphism of symbiotic genes encoding two components of the plant-bacteria signalling system. The first component is the rhizobialnodA acyltransferase involved in the fatty acid tail decoration of Nod factor (rhizobia signalling molecule). The second component is the plantNFR5receptor, putatively required for Nod-factor binding.We collected three wild growing legume species together with soil samples adjacent to the roots (soil pool) from one large 25-year fallow:Vicia sativa, Lathyrus pratensisandTrifolium hybridumnodulated by one of the twoRhizobium leguminosarumbiovars (viciaeandtrifolii). For each plant species we prepared three pools for DNA extraction: the plant pool (30 plant indiv.), the nodule pool (90 nodules) and the soil pool (30 samples).NFR5gene libraries from the plant pool andnodA gene libraries from nodule and soil pools were sequenced by Sanger technology and High-throughput pyrosequencing, respectively. Analysis of the data demonstrated concordance in population diversities of one symbiotic partner (rhizobia) the second partner (legume host), in line with the Evolutionary Moulding hypothesis. This effect was evinced by the following observations for each plant species: (1) significantly increased diversity in the nodulenodA popset (set of gene sequences derived from the nodule population) compared to the soil popset; (2) a monotonic relationship between the diversity in the plantNFR5gene popset and the nodule rhizobialnodA gene popset; and (3) higher topological similarity of theNFR5gene tree with thenodA gene tree of the nodule popset, than with thenodA gene tree of the soil popset. Both nonsynonymous diversity and Tajima’s D were increased in the nodule popsets compared to the soil popsets, consistent with relaxation of negative selection and/or admixture of balancing selection underlying the Evolutionary Moulding effect. We propose that the observed genetic concordance arises from the selection of particular characteristics of the nodulenodA genes by the host plant.