The members of the actinomycete genus Frankia are nitrogen-fixing symbionts of many species of woody dicotyledonous plants belonging to eight families. Several strains isolated from diverse actinorhizal plants growing in different geographical areas were used in this study. The phylogenetic relationships of these organisms and uncharacterized microsymbionts that are recalcitrant to isolation in pure culture were determined by comparing complete 16s ribosomal DNA sequences. The resulting phylogenetic tree revealed that there was greater diversity among the Alnus-infective strains than among the strains that infect other host plants. The four main subdivisions of the genus Frankia revealed by this phylogenetic analysis are (i) a very large group comprising Frankia alni and related organisms (including Alnus rugosa Sp+ microsymbionts that are seldom isolated in pure culture), to which Casuarina-infective strains, a Myrica nagi microsymbiont, and other effective Alnus-infective strains are related; (ii) unisolated microsymbionts of Dryas, Coriaria, and Dutisca species; (iii) Elaeagnus-infective strains; and (iv) "atypical" strains (a group which includes an Anus-infective, non-nitrogen-fixing strain). Taxa that are related to this well-defined, coherent Frankia cluster are the genera Geodermatophilus, "Blastococcus," Sporichthya, Acidothermus, and Actinoplanes. However, the two genera whose members have multilocular sporangia (the genera Frankia and Geodermatophilus) did not form a coherent group. For this reason, we propose that the family Frankiaceae should be emended so that the genera Geodermatophilus and "Blastococcus" are excluded and only the genus Frankia is retained.The slowly growing members of the actinomycete genus Frankia are root symbionts that nodulate a wide range of perennial woody dicotyledonous plants. This nitrogen-fixing symbiosis is known to occur in more than 200 species of plants belonging to 24 genera and eight families that are called actinorhizal (6). The first Frankia strain was isolated in 1956 by Pommer (43), but this strain was subsequently lost. In 1978, Callaham et al. (8) isolated an infective Frankia strain from Comptonia peregrirza, and since then hundreds of isolates have been obtained from a number of plant species growing in many geographical areas.Becking (5) was unsuccessful in isolating the causative agent of actinorhizal nodules despite numerous attempts. He suggested that this organism was an "obligate symbiont" and devised a classification scheme based on cross-inoculation groups and on the morphology of the endosymbiont. This scheme was subsequently found to be erroneous when pure cultures became available (8). Nonetheless, Becking ( 5 ) correctly perceived the bacterial nature of the microsymbiont, named it Frankia sp., and classified it as the only member of the family Frankiaceae in the order Actinomycetales.The members of the genus Frankia can now be clearly distinguished from other bacterial genera on the basis of their host specificity, their morphology (hyphae, ...
Some physical, thermal, and chemical properties of bark of 16 tree species native to the central hardwood region were measured to determine their potential to protect the vascular cambium from damage by fire. The relationship between DBH and bark thickness for each of 16 species was determined. For purposes of monitoring seasonal trends, two species (Quercusmacrocarpa Michx. and Acersaccharinum L.) were sampled periodically during one growing season. Temperature response to bark surface heating of 11 species was monitored at the cambial layer during simulated fires conducted in the field. Bark samples were analyzed for moisture content, specific gravity, dry weight, volatile matter content, and time until ignition. Overall, during simulated fires, temperature gradients were decreased and maximum cambial temperatures were reduced as bark thickness increased. Thick-barked species had lower maximum cambial temperatures, longer times to reach peak temperatures, slower rates of heat loss, and shorter time until surface ignition. Populusdeltoides Marsh, was the most heat resistant among species tested, while Acersaccharinum was the least. Higher specific gravities were associated with higher rates at which cambial temperatures rose as well as with increased time required for surface ignition.
Molecular phylogenetic approaches have begun to outline the origin, distribution and diversity of actinorhizal partners. Geographic isolation of Frankia and its host plants resulting from shifting continents and dispersal patterns have apparently led to the development of Frankia genotypes with differing affinities for host genera, even within the same plant family. Actinorhizal plant genera of widespread global distribution tend to nodulate readily even outside their native ranges. These taxa may maintain infective Frankia populations of considerable diversity on a broad scale. Arid environments seem to have distinctive actinorhizal partnerships, with smaller and more specific sets of Frankia symbionts. This has led to the hypothesis that some host families have taxa that are evolving towards narrow strain specificity, perhaps because of drier habitats where fewer Frankia strains would be able to survive. Harsh conditions such as water‐saturated soils near lakes, swamps or bogs that are typically acidic and low in oxygen may similarly lessen the diversity of Frankia strains present in the soil, perhaps limiting the pool of frankiae available for infection locally and, at a larger scale, for natural selection of symbiotic partnerships with host plants. Recent molecular ecological studies have also provided examples of Frankia strain sorting by soil environment within higher order cluster groupings of Frankia host specificity. Future frontiers for ecological research on Frankia and actinorhizal plants include the soil ecosystem and the genome of Frankia and its hosts.
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