Frankia alni | |
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Nodules caused by Frankia alni on roots of common alder Alnus glutinosa | |
Scientific classification | |
Domain: | Bacteria |
Phylum: | Actinomycetota |
Class: | Actinomycetia |
Order: | Frankiales |
Family: | Frankiaceae |
Genus: | Frankia |
Species: | F. alni |
Binomial name | |
Frankia alni (Woronin 1866) Von Tubeuf 1895 | |
Synonyms [1] [2] | |
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Frankia alni is a Gram-positive species of actinomycete filamentous bacterium that lives in symbiosis with actinorhizal plants in the genus Alnus . It is a nitrogen-fixing bacterium and forms nodules on the roots of alder trees.
Frankia alni forms a symbiotic relationship exclusively with trees in the genus Alnus. These are widely distributed in temperate regions of the northern hemisphere. One species, Alnus glutinosa , is also found in Africa and another, the Andean alder, Alnus acuminata , extends down the mountainous spine of Central and South America as far as Argentina. Evidence suggests that this alder may have been exploited by the Incas and used to increase soil fertility and stabilize terrace soils in their upland farming systems. [3] Actinomycetota, like Frankia alni, need a flagellum to be mobile, but F. alni does not have one, and is immobile. Alnus species grow in a wide range of habitats that include glacial till, sand hills, the banks of water courses, bogs, dry volcanic lava flows and ash alluvium. [4]
The first symptom of infection by Frankia alni is a branching and curling of the root hairs of the alder as the bacterium moves in. The bacterium becomes encapsulated with a material derived from the plant cell wall and remains outside the host's cell membrane. [5] The encapsulation membrane contains pectin, cellulose and hemicellulose. [6] Cell division is stimulated in the hypodermis and cortex, which leads to the formation of a "prenodule". The bacterium then migrates into the cortex of the root while the nodule continues to develop in the same way as a lateral root. Nodule lobe primordia develop in the pericycle, endodermis or cortex during the development of the prenodule and finally the bacterium enters the cells of these to infect the new nodule. [7]
In nitrogen-free culture and often in symbiosis, Frankia alni bacteria surround themselves in "vesicles". These are roughly spherical cellular structures that measure two to six millimetres in diameter and have a laminated lipid envelope. The vesicles serve to limit the diffusion of oxygen, thus assisting the reduction process that is catalysed by the enzyme nitrogenase. This enzyme bonds each atom of nitrogen to three hydrogen atoms, forming ammonia (NH3). The energy for the reaction is provided by the hydrolysis of Adenosine triphosphate (ATP). Two other enzymes are also involved in the process, glutamine synthetase and glutamate synthase. The final product of the reactions is glutamate, which is thus normally the most abundant free amino acid in the cell cytoplasm. A by-product of the process is gaseous hydrogen, one molecule of which is produced for every molecule of nitrogen reduced to ammonia, but the bacterium also contains the enzyme hydrogenase, which serves to prevent some of this energy being wasted. In the process, ATP is recovered and oxygen molecules serve as the final electron acceptor in the reaction, leading to the lowering of ambient oxygen levels. This is to the benefit of the nitrogenases, which only function anaerobically. [8]
As a result of their mutually beneficial relationship with Frankia, alder trees improve the fertility of the soils in which they grow and are considered to be a pioneer species, making the soil more fertile and thus enabling other successional species to become established.
In culture and in some root nodules, multilocular sporangia containing many spores are produced. [9] The sporangia are non-motile but the spores can migrate to infect new host plants. [10] A Swedish study found that root nodules developed on transplanted seedlings of the grey alder, Alnus incana, planted in meadow soil that had not grown actinorhizal plants for nearly sixty years. A similar experiment planting seedlings in deep layers of peat where the surface layer had been removed, did not produce nodulation. This seems to have been because there were no infective propagules of Frankia alni deep in the peat. No air-borne dispersal of Frankia alni was detected and it was thought that movement of water might account for the dispersal of the bacteria in peat soils. [11]
Alders are trees that compose the genus Alnus in the birch family Betulaceae. The genus includes about 35 species of monoecious trees and shrubs, a few reaching a large size, distributed throughout the north temperate zone with a few species extending into Central America, as well as the northern and southern Andes.
Nitrogen fixation is a chemical process by which molecular dinitrogen is converted into ammonia. It occurs both biologically and abiologically in chemical industries. Biological nitrogen fixation or diazotrophy is catalyzed by enzymes called nitrogenases. These enzyme complexes are encoded by the Nif genes and contain iron, often with a second metal.
Leghemoglobin is an oxygen-carrying phytoglobin found in the nitrogen-fixing root nodules of leguminous plants. It is produced by these plants in response to the roots being colonized by nitrogen-fixing bacteria, termed rhizobia, as part of the symbiotic interaction between plant and bacterium: roots not colonized by Rhizobium do not synthesise leghemoglobin. Leghemoglobin has close chemical and structural similarities to hemoglobin, and, like hemoglobin, is red in colour. It was originally thought that the heme prosthetic group for plant leghemoglobin was provided by the bacterial symbiont within symbiotic root nodules. However, subsequent work shows that the plant host strongly expresses heme biosynthesis genes within nodules, and that activation of those genes correlates with leghemoglobin gene expression in developing nodules.
Rhizobia are diazotrophic bacteria that fix nitrogen after becoming established inside the root nodules of legumes (Fabaceae). To express genes for nitrogen fixation, rhizobia require a plant host; they cannot independently fix nitrogen. In general, they are gram negative, motile, non-sporulating rods.
Alnus glutinosa, the common alder, black alder, European alder, European black alder, or just alder, is a species of tree in the family Betulaceae, native to most of Europe, southwest Asia and northern Africa. It thrives in wet locations where its association with the bacterium Frankia alni enables it to grow in poor quality soils. It is a medium-sized, short-lived tree growing to a height of up to 30 metres (98 feet). It has short-stalked rounded leaves and separate male and female flowers in the form of catkins. The small, rounded fruits are cone-like and the seeds are dispersed by wind and water.
Alnus rubra, the red alder, is a deciduous broadleaf tree native to western North America.
Dryas is a genus of perennial cushion-forming evergreen dwarf shrubs in the family Rosaceae, native to the arctic and alpine regions of Europe, Asia and North America. The genus is named after the dryads, the tree nymphs of ancient Greek mythology. The classification of Dryas within the Rosaceae has been unclear. The genus was formerly placed in the subfamily Rosoideae, but is now placed in subfamily Dryadoideae.
Diazotrophs are bacteria and archaea that fix atmospheric nitrogen (N2) in the atmosphere into bioavailable forms such as ammonia.
Root nodules are found on the roots of plants, primarily legumes, that form a symbiosis with nitrogen-fixing bacteria. Under nitrogen-limiting conditions, capable plants form a symbiotic relationship with a host-specific strain of bacteria known as rhizobia. This process has evolved multiple times within the legumes, as well as in other species found within the Rosid clade. Legume crops include beans, peas, and soybeans.
Nod factors, are signaling molecules produced by soil bacteria known as rhizobia in response to flavonoid exudation from plants under nitrogen limited conditions. Nod factors initiate the establishment of a symbiotic relationship between legumes and rhizobia by inducing nodulation. Nod factors produce the differentiation of plant tissue in root hairs into nodules where the bacteria reside and are able to fix nitrogen from the atmosphere for the plant in exchange for photosynthates and the appropriate environment for nitrogen fixation. One of the most important features provided by the plant in this symbiosis is the production of leghemoglobin, which maintains the oxygen concentration low and prevents the inhibition of nitrogenase activity.
Alnus incana, the grey alder, tag alder or speckled alder, is a species of multi-stemmed, shrubby tree in the birch family, with a wide range across the cooler parts of the Northern Hemisphere. Tolerant of wetter soils, it can slowly spread with runners and is a common sight in swamps and wetlands.
Frankia is a genus of nitrogen-fixing bacteria that live in symbiosis with actinorhizal plants, similar to the Rhizobium bacteria found in the root nodules of legumes in the family Fabaceae. Frankia also initiate the forming of root nodules.
Alnus cordata, the Italian alder, is a tree or shrub species belonging to the family Betulaceae, and native to the southern Apennine Mountains and the north-eastern mountains of Corsica. It has been introduced in Sicily, Sardinia, and more recently in Central-Northern Italy, other European countries and extra-European countries, where it has become naturalised.
Chamaebatia, also known as mountain misery, is a genus of two species of aromatic evergreen shrubs endemic to California. Its English common name derives from early settlers' experience with the plant's dense tangle and sticky, strong-smelling resin. They are actinorhizal, non-legumes capable of nitrogen fixation through symbiosis with the actinobacterium, Frankia.
Phytophthora alni is an oomycete plant pathogen that causes lethal root and collar rot in alders. It is widespread across Europe and has recently been found in North America. This species is believed to have originated relatively recently.
The nif genes are genes encoding enzymes involved in the fixation of atmospheric nitrogen into a form of nitrogen available to living organisms. The primary enzyme encoded by the nif genes is the nitrogenase complex which is in charge of converting atmospheric nitrogen (N2) to other nitrogen forms such as ammonia which the organism can use for various purposes. Besides the nitrogenase enzyme, the nif genes also encode a number of regulatory proteins involved in nitrogen fixation. The nif genes are found in both free-living nitrogen-fixing bacteria and in symbiotic bacteria associated with various plants. The expression of the nif genes is induced as a response to low concentrations of fixed nitrogen and oxygen concentrations (the low oxygen concentrations are actively maintained in the root environment of host plants). The first Rhizobium genes for nitrogen fixation (nif) and for nodulation (nod) were cloned in the early 1980s by Gary Ruvkun and Sharon R. Long in Frederick M. Ausubel's laboratory.
Actinorhizal plants are a group of angiosperms characterized by their ability to form a symbiosis with the nitrogen fixing actinomycetota Frankia. This association leads to the formation of nitrogen-fixing root nodules.
The flavans are benzopyran derivatives that use the 2-phenyl-3,4-dihydro-2H-chromene skeleton. They may be found in plants. These compounds include the flavan-3-ols, flavan-4-ols and flavan-3,4-diols (leucoanthocyanidin).
A nitrogen fixation package is a piece of research equipment for studying nitrogen fixation in plants. One product of this kind, the Q-Box NF1LP made by Qubit Systems, operates by measuring the hydrogen (H2) given off in the nitrogen-fixing chemical reaction enabled by nitrogenase enzymes.
A symbiosome is a specialised compartment in a host cell that houses an endosymbiont in a symbiotic relationship.