Determinate development of nodule roots in actinomycete-induced root nodules of Myrica gale

Abstract

Young seedlings of Myrica gale L. grown in water culture were inoculated with a nodule suspension containing the effective actinomycete which induced root nodule formation. Nodule development was followed from initiation to nodule lobe formation and nodule root development using living materials and fixed nodules sectioned for light microscopy. After root hair infection and prenodule formation, three stages were observed: nodule lobe formation, a transition or arrested state, and nodule root development. The primary nodule lobe meristem originates endogenously and its formation involves pericycle, endodermis, and cortical cell derivatives. The lobe develops slowly to about 2 mm in length while the cortical cells are invaded by the actinomycete endophyte. After a period of arrest of variable duration, from a few days to several weeks, the nodule lobe meristem begins altered development, forming the elongate nodule root which undergoes slow but continuous growth to about 3- to 4-cm final length. New nodule lobe primordia are initiated endogenously at the base of existing nodules lobes, ultimately forming a cluster of nodule roots. Each nodule root, which elongates at about 0.1–1.0 mm per day, has a terminal apical meristem with reduced root cap formation and produces a modified root structure possessing an elaborate cortical intercellular space system and a reduced central cylinder. Nodule root growth is distinctive in that it shows strong negative geotropism. The endophyte is restricted to cortical cells of the nodule lobe and is totally absent from tissues of the nodule root. A probable role for nodule roots is to facilitate gas diffusion to the nitrogen-fixing endophyte site in the nodule lobe when nodules occur under conditions of low oxygen tension.