Haustorium
Encyclopedia
In botany, a haustorium (plural haustoria) is the appendage or portion of a parasitic
Parasitism
Parasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host . These are now called macroparasites...

 fungus (the hyphal tip) or of the root of a parasitic plant
Parasitic plant
A parasitic plant is one that derives some or all of its sustenance from another plant. About 4,100 species in approximately 19 families of flowering plants are known. Parasitic plants have a modified root, the haustorium, that penetrates the host plant and connects to the xylem, phloem, or...

 (such as the broomrape family
Orobanchaceae
Orobanchaceae, the broomrape family, is a family of flowering plants of the order Lamiales, with about 90 genera and more than 2000 species. Many of these genera were formerly included in the family Scrophulariaceae sensu lato...

 or mistletoe
Mistletoe
Mistletoe is the common name for obligate hemi-parasitic plants in several families in the order Santalales. The plants in question grow attached to and within the branches of a tree or shrub.-Mistletoe in the genus Viscum:...

) that penetrates the host's tissue and draws nutrients from it. Haustoria do not penetrate the host's cell membranes.

Fungi in all major divisions form haustoria. Haustoria take several forms. Generally, on penetration, the fungus increases the surface area in contact with host plasma membrane releasing enzymes that break down the cell wall, enabling greater potential movement of organic carbon from host to fungus. Thus, an insect hosting a parasitic fungus such as Cordyceps
Cordyceps
Cordyceps is a genus of ascomycete fungi that includes about 400 described species. All Cordyceps species are endoparasitoids, mainly on insects and other arthropods ; a few are parasitic on other fungi. The best known species of the genus is Cordyceps sinensis, first recorded as yartsa gunbu in...

 may look as though it is being "eaten from the inside out" as the haustoria expand inside of it.
Haustoria arise from intercellular hyphae, appressoria, or external hyphae. The hypha narrows as it passes through the wall
Cell wall
The cell wall is the tough, usually flexible but sometimes fairly rigid layer that surrounds some types of cells. It is located outside the cell membrane and provides these cells with structural support and protection, and also acts as a filtering mechanism. A major function of the cell wall is to...

 of the cell and then expands on invaginating the cell. A thickened, electron-dense collar of material is deposited around the hypha at the point of invagination. Further, the host wall becomes highly modified in the invaginated zone. Inclusions normally present in plasma membrane are absent, and the outer layer contains more polysaccharide. The wall of both partners is severely reduced.

Functional exchange takes place within the haustorial complex. The host supplies organic carbon
Carbon
Carbon is the chemical element with symbol C and atomic number 6. As a member of group 14 on the periodic table, it is nonmetallic and tetravalent—making four electrons available to form covalent chemical bonds...

 to the fungus, and the metabolic activity within the complex is considerably greater than outside. Carbon from the host is absorbed by the fungus, and immediately transported to the rest of the thallus. The host plant appears to be functioning according to signals from the fungus and the complex appears to be under the control of the invader.

The haustorium may be balloon-, spiral- or glove-shaped.
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