Alders (Alnus spp.) are important components of northern ecosystems due to their ability to fix nitrogen (N) in symbiosis with Frankia bacteria. Availability of optimal Frankia may be a contributing factor in limiting the performance and ecological effects of Alnus, but the factors underlying distribution of Alnus-infective Frankia are not well understood. This study examined the genetic structure (nifD-K spacer RFLP haplotypes) of Frankia assemblages symbiotic with two species of Alnus (A. tenuifolia and A. viridis) in four successional habitats in interior Alaska. We used one habitat in which both hosts occurred to observe differences between host species independent of habitat, and we used replicate sites for each habitat and host to assess the consistency of symbiont structure related to both factors. We also measured leaf N content and specific N-fixation rate (SNF) of nodules ((15)N uptake) to determine whether either covaried with Frankia structure, and whether Frankia genotypes differed in SNF in situ. Frankia structure differed between sympatric hosts and among habitats, particularly for A. tenuifolia, and was largely consistent among replicate sites representing both factors. Leaf N differed between host species and among habitats for both hosts. SNF did not differ among habitats or host species, and little evidence for differences in SNF among Frankia genotypes was found, due largely to high variation in SNF. Consistency of Frankia structure among replicate sites suggests a consistent relationship between both host species and habitat among these sites. Correlations with specific environmental variables and possible underlying mechanisms are discussed.
In long-lived N-fixing plants, environmental conditions affecting plant growth and N demand vary at multiple temporal and spatial scales, and symbiont assemblages on a given host and patterns of allocation to nodule activities have been shown to vary according to environmental factors, suggesting that hosts may alter partner choice and manipulate symbiont assemblages based on shifting plant needs. This study assessed economic trade-offs among N-fixing symbionts of thin-leaf alder (Alnus tenuifolia) by examining whether alderFrankia associations change in response to the up-regulation (P fertilization) and downregulation (N-fertilization) of N-fixation activities, and whether these changes are associated with differences among Frankia partners in their relative C cost and/or N benefit to A. tenuifolia.Relative to control plots, alder in þP plots had significantly higher nodule biomass and Nfixation rates; these parameters were significantly lower in þN plots, translating to stand-level N inputs that were more than an order of magnitude greater in þP than þN plots. Nodule respiration and N-fixation rates were positively correlated, and analyses revealed that alder employs mechanisms to increase the efficiency of C use when N fixation is up-regulated. Of the eight Frankia OTUs (operational taxonomic units) identified, two were dominant, with significant differences in Frankia OTU composition across samples being explained by fertilization treatment. Dominant OTUs had similar up-and down-regulatory responses to treatments but differed in C costs of N fixation, while the most abundant sub-dominant failed to up-regulate N-fixation rates in þP plots. Differences among Frankia OTUs in traits relevant to plant performance may form the basis for host choice and explain successional shifts in alder-Frankia assemblages. We suggest that manipulation of Frankia assemblages is an adaptation for a species such as A. tenuifolia that maintains a high dependence on atmospheric N across radically different growth environments. Because of the strong effects of thin-leaf alder on soil N contents, it appears that host changes in C partitioning to nodule activities, including the physiological consequences of host specificity for Frankia assemblages, results in alder both driving and responding to environmental heterogeneity at small to large spatial scales.
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