How are nitrogen fixation rates regulated in legumes?

Symbiotic nitrogen fixation by the legume-Rhizobium symbiosis is a finely regulated process that involves significant carbon and energy metabolism by the plant. At present, there are three competing theories as to how the regulation of the process is achieved at the whole plant level. Theory one (carbon supply regulation) states that nitrogen fixation rates are regulated by either current nodule assimilate supply or through metabolism of assimilates into compounds that can be used by the bacteroids. A second theory (oxygen supply regulation) assumes that oxygen diffusion into the nodules is tightly regulated and is the principal regulatory factor for nitrogen fixation rates. A third theory (N-feedback regulation) suggests that a product of nitrogen fixation or assimilation exerts a feedback regulatory impact. The paper summarizes experimental data which support or reject the mentioned theories. Moreover, implications of nitrogen fixation under P stress and of phenomena connected with the argon or acetylene induced decline in nitrogenase activity for the understanding of nitrogen fixation regulation are discussed. It is concluded that we currently have no theory that explains all aspects and experimental results concerning the regulation of nitrogen fixation in legumes. With our growing ability to impact the process, e.g., by genetic engineering, this understanding of nitrogen fixation regulation has potential to be translated into agronomically sustainable benefits.