Nitrogen Fixation by Nodulated Roots of Viminaria juncea (Schrad. & Wendl.) Hoffmans, (Fabaceae) When Submerged in Water

Abstract

Nodulated seedlings of Viminaria juncea were raised in free-draining or flooded sand culture. Unflooded seedlings developed limited amounts of aerenchyma in lower stem, root and nodules, and responded to flooding by accelerated aerenchyma production and, after 10 days, by formation of pneumatophores from their near-surface lateral roots. Continuously flooded seedlings showed earlier and greater development of aerenchyma and pneumatophores, and had their nodules and roots restricted to the upper 10 cm of the rooting medium. Aerenchyma was developed from an inner cambium, distinct from the outer phellogen which subsequently developed on older parts of stem, root and nodules. Gas contents of plant parts varied from 4-8% for organs with little aerenchyma to over 30% for the aerenchyma-invested basal stem and root of continuously flooded seedlings. A role of the sheaths of aerenchyma in gaseous exchange between aerial environment and nodulated root was demonstrated by gas injection experiments, in situ C2H2 reduction assays and 15N2 feeding experiments on intact plants with flooded roots. Samples of gas removed from the aerenchyma of plants exposed to C2H2 contained up to 14 times the amount of C2H2 and 4 times the amount of CO2 than in the atmosphere of the assay chamber, indicating that gas exchange for both N2 fixation and respiration occurred via the aerenchyma. Previously unflooded, 12-week seedlings exposed to 14 days flooding gained as much dry matter and total N in the 2-week treatment as did control unflooded plants, but 21-week continuously flooded seedlings showed only half the dry matter and nitrogen gains of similarly aged unflooded seedlings. Observations on the seasonal growth, nodulation and pneumatophore development of natural populations of the species were discussed in relation to the above findings.