Frankia is a genus of soil actinomycetes famous for its ability to form N 2 -fixing root nodule symbioses with actinorhizal plants. Although Frankia strains display a high diversity in terms of ecological niches in soil, current knowledge about Frankia is dominated by its life as an endophyte in root nodules. Increased use of molecular methods has refined and expanded insights into endophytehost specificities and Frankia phylogeny. This review has focus on Frankia as a soil organism, including its part of microbial consortia, and how to study Frankia in soil. We highlight the use of nodulation tests and molecular methods to reveal population size and genetic diversity of Frankia in soil and discuss how autoregulation of nodulation and interactions with other soil microorganisms may influence the results. A comprehensive record of published interactions between Frankia and other soil microbes is summarized. Life in soil by the actinorhizal root nodule endophyte Frankia 203 204 E.E. Chaia et al. E.E. Chaia et al.

Leys, used for grazing or production of forage to be conserved as silage or hay, are very important crops in northern areas. In order to measure the N 2 fixation in leys of varying ages and during different parts of the season, detailed measurements were taken of yield, N 2 fixation and the amounts of N remaining in the field after harvesting red clover (Trifolium pratense L.)-grass leys at a site in northern Sweden, where they are generally harvested twice per growing season. Entire plants, including stubble and roots, were sampled at the time of first and second harvest and, in addition, at the end of the growing season in three neighbouring fields, carrying a first, a second and a third year ley, respectively. N 2 fixation was measured by both 15 N isotope dilution (ID) and 15 N natural abundance (NA) methods. The proportion of clover dry matter (DM) in the stands increased from the first to the second harvest, but the grasses dominated throughout the entire season, especially below ground. The N concentrations, in both herbage and whole plants, were about twice as high in the clover as in the grasses. Seasonal variations in N concentrations were minor, and total N contents followed the same trends as DM. The clover acquired nearly all of its N from N 2 fixation: the proportion of N in clover herbage derived from N 2 fixation was often >0.8 throughout the season. The variations in the amounts of N 2 fixed during the course of the season corresponded well to the seasonal changes in clover biomass. Amounts of fixed N 2 allocated to clover herbage during the whole season were in the range 4 to 6 g N m −2 in this unusually rainy year. Calculations of daily N allocation rates to herbage showed that N uptake rates were similar, and high, in grasses during May-June and July-August, while N 2 fixation rates in clover were about 10-fold as high in July-August as in May-June, reflecting the need for N in clover growth. The proportion of N remaining in clover stubble and roots after the first and second harvests was about 60 and 25%, respectively, while about 60% of the N in grasses remained in stubble and roots after both harvests. The considerable amounts of biomass and N that were left in field after harvesting red clover-grass leys are important for regrowth of the plants and provide substantial N fertilization for the next crop in the crop rotation.

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The infective capacities of the nitrogen fixing Actinomycete Frankia and arbuscular mycorrhizal fungi from soils near watercourses, along a vegetation gradient, were studied using plant bioassays. Frankia and arbuscular mycorrhizas capable of infecting Discaria trinervis were found at seventeen sites sampled. More specific enumeration of the infective capacities of both microorganisms in relation to environmental factors was performed in seven representative soils of the analysed vegetation zones (rainforest, xeric forest and steppe) using the most probable number method. The highest nodulation capacities ranged from 340 infective units g(-1 )soil, in a steppe marsh devoid of actinorhizas, to 61 in a coastal actinorhizal scrub (in xeric forest). The highest number of infective mycorrhizal units--also found in marsh--was 145. In general, rainforest soils had the lowest values for both microorganisms. Infective units of Frankia and arbuscular mycorrhizal fungi in soil were positively correlated (r = 0.89, P < 0.05). Both soilborne symbionts showed the highest infective capacity in semi-arid conditions nearby watercourse and at the valley bottom location. Tripartite symbiosis was effective in plants inoculated with steppe and xeric forest soils and plants inoculated with Frankia BCU110501 and Glomus mosseae. Interaction between both symbionts and influence of environmental conditions, in general, would contribute to define comparable trends of their infective capacities.