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Index >> Rhizobium and Legume Root Nodulation >>Structure of the Nodule

Structure of the Nodule

Structure of the Nodule
The core of a mature nodule constitutes the 'bacteroid zone' which is surrounded by several layers of cortical cells. The relative volume of bacteroid tissue (16 to 50% of the dry weight of the nodules) is much greater in effective nodules than in ineffective ones. The volume of bacteroid tissue in effective nodules has a direct positive relationship with the amount of nitrogen fixed. Ineffective nodules produced by ineffective strains are generally small and contain poorly developed bacteroid tissue associated with structural abnormalities.

In all ineffective associations, it has been shown that starch accumulates in the uninfected cell and dextran in the infected cell with glycogen in the bacteroid. Effective nodules are generally large and pink (due to leghaemoglobin) with well developed and organized bacteroid tissue.

A fully-developed bacteroid has no flagella and is surrounded by three unit membranes. There exists an intracytoplasmic membrane system in the bacteroids of nodules in subterranean clover. The nuclear region of bacteroids appears fragmented and is associated with granular cytoplasm. Bacteroids can be produced in vitro on a medium containing 3.5% of yeast extract. Caffeine and several other alkaloids also encourage bacteroid production on artificial media. Depending on the legume, each bacteroid or groups of bacteroids are surrounded by membrane envelopes whose identity has been variously interpreted, possibly because of the different techniques used in fine structure studies.

The three hypotheses concerning them are: (1) they are formed de novo after the release of the bacteria from the infection thread, (2) they are extensions of the endoplasmic reticulum of the host cells, and (3) they are derived from plasmalemma by a process of phagocytosis. The number of bacteroids enclosed in membrane envelopes appears to vary from one to many depending on the species of legume.

The membrane envelope surrounding the bacteroids is also known as the peribacteroid membrane. The earlier concept that the peribacteroid membrane is only a slightly modified plasma membrane has to be viewed differently in the light of later findings. The multiplication of bacteroids and the formation of the peribacteroid membrane may not be synchronized cell events which result in the variability in the number of bacteroids enclosed by the membrane. The microsymbiont (Rhizobium) strain determines the particle density and the protein and fatty acid composition of the membrane. The peribacteroid membrane contains nodulins which may play special roles in the two-way transport of metabolites between the symbionts.

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