Root nodule

Root nodules occur on the roots of plants that associate with symbiotic bacteria.

Under nitrogen limiting conditions, plants from the pea family Fabaceae form a symbiotic relationship with a host-specific strain of bacteria known as rhizobia.

Within legume nodules, nitrogen gas from the atmosphere is converted into ammonia, which then is assimilated by the plant to form the basis for amino acids (the building blocks of proteins), nucleotides (the building blocks of DNA and RNA as well as the important energy molecule ATP), and other cellular constituents such as vitamins, flavones, and hormones. The nitrogen fixation property makes legumes an ideal agricultural organism as their requirement for nitrogen fertiliser is reduced. Indeed high nitrogen content blocks nodule development. The energy for splitting the nitrogen gas in the nodule comes from sugar that is translocated from the leaf (a product of photosynthesis). Malate as a breakdown product of sucrose is the direct carbon source for the bacteroid. Nitrogen fixation in the nodule is very oxygen sensitive. Legume nodules harbor an iron containing protein called hemoglobin, closely related to animal myoglobin, to facilitate the conversion of nitrogen gas to ammonia.

Classification
Two main types of nodule have been described.

Temperate legumes like Pisum, Medicago, Trifolium, and Vicia develop a cylindrical shaped nodule that is called "indeterminate" because it maintains an active apical meristem that produces new cells for growth over the life of the nodule. The genus Lupinus is nodulated by the soil microorganism [http://lupins-bk.blogspot.com/2006/07/nitrogen-fixation.html Bradyrhizobium sp. (Lupinus).] Bradyrhizobia are encountered as microsymbionts in other leguminous crops (Argyrolobium, Lotus, Ornithopus, Acacia, Lupinus) of Mediterranean origin

Tropical (sub)legumes from the genera Glycine, Phaseolus, Lotus, and Vigna form "determinate" nodules, that lose meristematic activity shortly after initiation. Growth is due to cell expansion, and mature nodules are spherical in shape.

Nodulation
Legumes release compounds called flavonoids from their roots, which trigger the production of nod factors by the bacteria. When the nod factor is sensed by the root, a number of biochemical and morphological changes happen: cell division is triggered in the root to create the nodule, and the root hair growth is redirected to wind around the bacteria multiple times until it fully encapsulates 1 or more bacteria. The bacteria encapsulated divide multiple times, forming a microcolony. From this microcolony, the bacteria enter the developing nodule through a structure called an infection thread, which grows through the root hair into the basal part of the epidermis cell, and onwards into the root cortex; they are then surrounded by a plant-derived membrane and differentiate into bacteroids that fix nitrogen.

Nodulation is controlled by a variety of processes, both external (heat, acidic soils, drought, nitrate) and internal (autoregulation of nodulation, ethylene). Autoregulation of nodulation controls nodule numbers per plant thorugh a systemic process involving the leaf. Leaf tissue senses the early nodulation events in the root through an unknown chemical signal, then restricts further nodule development in newly develing root tissue. The Leucine rich repeat (LRR) receptor kinases (NARK in soybean (Glycine max); HAR1 in Lotus japonicus, SUNN in Medicago truncatula) are essential for autoregulation of nodulation (AON). Mutation leading to loss of function in these AON receptor kinases leads to supernodulation or hypernodulation. Often root growth abnormalities accompany the loss of AON receptor kinase activity, suggesting that nodule growth and root development are functionally linked.

In other species
Root nodules that occur on non-legume genera like Parasponia in association with Rhizobium bacteria, and those that arise from symbiotic interactions with Actinobacteria Frankia in some plant genera such as Alnus, vary significantly from those formed in the legume-rhizobia symbiosis. In these symbioses the bacteria are never released from the infection thread.

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