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Soil bacteria, collectively called rhizobia    , symbiotically interact with legume roots to form specialized structures called nodules    , in which nitrogen fixation takes place. This process entails the reduction of atmospheric nitrogen to ammonia, by means of the enzyme nitrogenase    . Therefore, using rhizobia is a natural and environmentally friendly way to fertilize plants, as opposed to chemical fertilization that uses a nonrenewable resource, such as natural gas. Through symbiotic nitrogen fixation, the plant benefits from using an endless source of nitrogen from the atmosphere. The process simultaneously contributes to soil fertility because the plant root system leaves behind some of the biologically available nitrogen. As in any symbiosis, both organisms benefit from the interaction: the plant obtains ammonia, and bacteria obtain carbon compounds generated through photosynthesis, as well as a protected niche in which to grow ( [link] ).

 Part A is a photo of legume roots, which are long and thin with hair-like appendages. Nodules are bulbous protrusions extending from the root. Part B is a transmission electron micrograph of a nodule cell cross section. Black oval-shaped vesicles containing rhizobia are visible. The vesicles are surrounded by a white layer and are scattered unevenly throughout the cell, which is gray.
Soybean roots contain (a) nitrogen-fixing nodules. Cells within the nodules are infected with Bradyrhyzobium japonicum, a rhizobia or “root-loving” bacterium. The bacteria are encased in (b) vesicles inside the cell, as can be seen in this transmission electron micrograph. (credit a: modification of work by USDA; credit b: modification of work by Louisa Howard, Dartmouth Electron Microscope Facility; scale-bar data from Matt Russell)

Mycorrhizae: the symbiotic relationship between fungi and roots

A nutrient depletion zone can develop when there is rapid soil solution uptake, low nutrient concentration, low diffusion rate, or low soil moisture. These conditions are very common; therefore, most plants rely on fungi to facilitate the uptake of minerals from the soil. Fungi form symbiotic associations called mycorrhizae with plant roots, in which the fungi actually are integrated into the physical structure of the root. The fungi colonize the living root tissue during active plant growth.

Through mycorrhization, the plant obtains mainly phosphate and other minerals, such as zinc and copper, from the soil. The fungus obtains nutrients, such as sugars, from the plant root ( [link] ). Mycorrhizae help increase the surface area of the plant root system because hyphae, which are narrow, can spread beyond the nutrient depletion zone. Hyphae can grow into small soil pores that allow access to phosphorus that would otherwise be unavailable to the plant. The beneficial effect on the plant is best observed in poor soils. The benefit to fungi is that they can obtain up to 20 percent of the total carbon accessed by plants. Mycorrhizae functions as a physical barrier to pathogens. It also provides an induction of generalized host defense mechanisms, and sometimes involves production of antibiotic compounds by the fungi.

 Photo shows a root with many branching tips. The surface of the root is fuzzy in appearance.
Root tips proliferate in the presence of mycorrhizal infection, which appears as off-white fuzz in this image. (credit: modification of work by Nilsson et al., BMC Bioinformatics 2005)

There are two types of mycorrhizae: ectomycorrhizae and endomycorrhizae. Ectomycorrhizae form an extensive dense sheath around the roots, called a mantle. Hyphae from the fungi extend from the mantle into the soil, which increases the surface area for water and mineral absorption. This type of mycorrhizae is found in forest trees, especially conifers, birches, and oaks. Endomycorrhizae, also called arbuscular mycorrhizae, do not form a dense sheath over the root. Instead, the fungal mycelium is embedded within the root tissue. Endomycorrhizae are found in the roots of more than 80 percent of terrestrial plants.

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Source:  OpenStax, Biology. OpenStax CNX. Feb 29, 2016 Download for free at http://cnx.org/content/col11448/1.10
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