Alnus maritima (Marsh.) Muhl. ex Nutt. is unique among alders in its degree of preference for low-oxygen soils of wetlands. An actinorhizal species with promise for use in sustainable horticulture, A. maritima develops a root-nodule symbiosis with nitrogen-fixing Frankia. Nodules of other actinorhizal species that are obligate wetland natives are adapted to low oxygen, and expression of hemoglobin is common to these taxa. Our objectives were to determine the range of oxygen tension under which Alnus maritima subsp. maritima fixes nitrogen and to investigate a potential role for hemoglobin in adaptation of nodules to low oxygen. Roots of plants, cultured aeroponically, were subjected to eight oxygen tensions from 0 to 32 kPa. After four weeks, plant dry weight, nodule fresh weight, nitrogenase activity, and photosynthetic rate were measured. In addition, nodules were assayed spectrophotometrically for the presence of hemoglobin. A quadratic function best described the influence of oxygen on plant dry weight, nodule fresh weight, nitrogenase activity, and photosynthetic rate with maximal values above 20 kPa. Alnus serrulata (Ait.) Willd. is sympatric with A. maritima subsp. maritima but is not an obligate inhabitant of wetlands. In a separate experiment, we found higher nitrogenase activity in A. maritima subsp. maritima than in A. serrulata (0.74 vs. 0.26 μmol/h per plant) at hypoxic oxygen tensions. Further, optical absorption spectra of nodule extracts confirmed hemoglobin within nodules of A. maritima subsp. maritima. Our data suggest that hemoglobin contributes to oxygen regulation in nodules of A. maritima subsp. maritima.
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