Development of flood-induced lenticels in red alder nodules prior to the restoration of nitrogenase activity
Red alder (Alnus rubra Bong.) is a nitrogen-fixing woody plant that is common on wetland sites and tolerates flooding through a variety of induced morphological and physiological changes. Among these changes are the formation of hypertrophied nodule lenticels and the subsequent full restoration of nitrogenase activity after 50 days of flooding. The objective of this study was to examine fine structural changes within red alder nodules during lenticel development that correspond to changes in nitrogenase activity during a 50-day experimental flood. Nodulated seedlings of red alder were grown under greenhouse conditions and then exposed to root flooding for 1, 20, 35, or 50 days. At each harvest, estimates of nitrogenase activity were made via acetylene reduction, and nodule samples were taken for light-microscope examination. Only after 50 days of flooding did red alder show restoration of nitrogenase to pretreatment levels. At this time, Frankia vesicles were found to be directly adjacent to developing lenticel tissue and large intercellular spaces. Intercellular space within the nodule increased from 0.6% in nonflooded tissue to 5.7% after 50 days of flooding. Our results demonstrate the sensitivity of the nitrogenase enzyme to low oxygen soil conditions and indicate that substantial morphological change within the nodule must occur before red alder can regain the capacity to fix nitrogen under flooded conditions.Key words: Alnus rubra, Frankia, waterlogged, biological nitrogen fixation, hypertrophied lenticels, intercellular space.