Mycorrhiza

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This mycorrhiza includes a fungus of the genus Amanita

A mycorrhiza (Greek: μυκός, mykós, "fungus" and ριζα, riza, "roots",[1] pl. mycorrhizae or mycorrhizas) is a symbiotic (generally mutualistic, but occasionally weakly pathogenic) association between a fungus and the roots of a vascular plant.[2]

In a mycorrhizal association, the fungus colonizes the host plant's roots, either intracellularly as in arbuscular mycorrhizal fungi (AMF or AM), or extracellularly as in ectomycorrhizal fungi. They are an important component of soil life and soil chemistry.

Mutualist dynamics[edit]

Mycorrhizae form a mutualistic relationship with the roots of most plant species. While only a small proportion of all species has been examined, 95% of those plant families[which?] are predominantly mycorrhizal.[3] They are named after their presence in the plant's rhizosphere (root system).

Recent research with ectomycorrhizal in boreal forests, has indicated mycorrhizae and plants have a relationship that may be more complex than simply mutualistic. This relationship was noted when mycorrhizae were unexpectedly found hoarding nitrogen from plant roots in times of nitrogen scarcity. Researchers argue that some mycorrhizae distribute nutrients based upon the environment with surrounding plants and other mycorrhizae, in a manner similar to a capitalistic market. Researchers go on to explain how this updated model explains why mycorrhizae do not alleviate plant nitrogen limitation, and why plants can switch abruptly from a mixed strategy with both mycorrhizal and nonmycorrhizal roots to a purely mycorrhizal strategy as soil nitrogen availability declines.[4]

Sugar-water/mineral exchange[edit]

This mutualistic association provides the fungus with relatively constant and direct access to carbohydrates, such as glucose and sucrose.[5] The carbohydrates are translocated from their source (usually leaves) to root tissue and on to the plant's fungal partners. In return, the plant gains the benefits of the mycelium's higher absorptive capacity for water and mineral nutrients due to the comparatively large surface area of mycelium: root ratio, thus improving the plant's mineral absorption capabilities.[6]

Plant roots alone may be incapable of taking up phosphate ions that are demineralized in soils with a basic pH. The mycelium of the mycorrhizal fungus can, however, access these phosphorus sources, and make them available to the plants they colonize.[7] Thus many plants are able to obtain phosphate, without using soil as a source. For example, in some dystrophic forests, large amounts of phosphate are taken up by mycorrhizal hyphae acting directly on leaf litter, bypassing the need for soil uptake.[8] Inga alley cropping, proposed as an alternative to slash and burn rainforest destruction,[9] relies upon mycorrhiza within the Inga Tree root system to prevent the rain from washing phosphorus out of the soil.[10] In some cases, the transport of water, carbon, and nutrients could be done directly from plant to plant through mycorrhizal networks that are underground hyphal networks created by mycorrhizal fungi that connect individual plants together.[11]

Suillus tomentosus, a basidiomycete fungus, produces specialized structures known as tuberculate ectomycorrhizae with its plant host lodgepole pine (Pinus contorta var. latifolia). These structures have been shown to host nitrogen fixing bacteria which contribute a significant amount of nitrogen and allow the pines to colonize nutrient-poor sites.[12]

Mechanisms[edit]

The mechanisms of increased absorption are both physical and chemical. Mycorrhizal mycelia are much smaller in diameter than the smallest root, and thus can explore a greater volume of soil, providing a larger surface area for absorption. Also, the cell membrane chemistry of fungi is different from that of plants (including organic acid excretion which aids in ion displacement[13]). Mycorrhizas are especially beneficial for the plant partner in nutrient-poor soils.[14]

Disease and drought resistance and its correlation to mycorrhizae[edit]

Mycorrhizal plants are often more resistant to diseases, such as those caused by microbial soil-borne pathogens. AMF was also significantly correlated with soil biological fertility variables such as soil fungi and soil bacteria, including soil disease.[citation needed] Furthermore, AMF was significantly correlated with soil physical variable, but only with water level and not with aggregate stability.[15][16] and are also more resistant to the effects of drought.[17][18][19]

Colonization of barren soil[edit]

Plants grown in sterile soils and growth media often perform poorly without the addition of spores or hyphae of mycorrhizal fungi to colonise the plant roots and aid in the uptake of soil mineral nutrients.[20] The absence of mycorrhizal fungi can also slow plant growth in early succession or on degraded landscapes.[21] The introduction of alien mycorrhizal plants to nutrient-deficient ecosystems puts indigenous non-mycorrhizal plants at a competitive disadvantage.[22]

Resistance to toxicity[edit]

Fungi have been found to have a protective role for plants rooted in soils with high metal concentrations, such as acidic and contaminated soils. Pine trees inoculated with Pisolithus tinctorius planted in several contaminated sites displayed high tolerance to the prevailing contaminant, survivorship and growth.[citation needed] One study discovered the existence of Suillus luteus strains with varying tolerance of zinc. Another study discovered that zinc-tolerant strains of Suillus bovinus conferred resistance to plants of Pinus sylvestris. This was probably due to binding of the metal to the extramatricial mycelium of the fungus, without affecting the exchange of beneficial substances.[22]

Occurrence of mycorrhizal associations[edit]

At around 400 million years old, the Rhynie chert contains the earliest fossil assemblage yielding plants preserved in sufficient detail to detect mycorrhizas - and they are indeed observed in the stems of Aglaophyton major.[23]

Mycorrhizas are present in 92% of plant families studied (80% of species),[24] with arbuscular mycorrhizas being the ancestral and predominant form,[24] and indeed the most prevalent symbiotic association found in the plant kingdom.[5] The structure of arbuscular mycorrhizas has been highly conserved since their first appearance in the fossil record,[23] with both the development of ectomycorrhizas, and the loss of mycorrhizas, evolving convergently on multiple occasions.[24]

Types of mycorrhiza[edit]

Arbuscular mycorrhizal wheat

Mycorrhizas are commonly divided into ectomycorrhizas and endomycorrhizas. The two types are differentiated by the fact that the hyphae of ectomycorrhizal fungi do not penetrate individual cells within the root, while the hyphae of endomycorrhizal fungi penetrate the cell wall and invaginate the cell membrane. Additionally, many plants in the order Ericales form a third type, ericoid mycorrhizas, while some members of the Ericales form arbutoid and monotropoid mycorrhizas.[25][26] All orchids are myco-heterotrophic at some stage during their lifecycle and form orchid mycorrhizas with a range of basidiomycete fungi.

Endomycorrhiza[edit]

Main article: Arbuscular mycorrhiza

Endomycorrhizas are variable and have been further classified as arbuscular, ericoid, arbutoid, monotropoid, and orchid mycorrhizas.[27] Arbuscular mycorrhizas, or AM (formerly known as vesicular-arbuscular mycorrhizas, or VAM), are mycorrhizas whose hyphae enter into the plant cells, producing structures that are either balloon-like (vesicles) or dichotomously branching invaginations (arbuscules). The fungal hyphae do not in fact penetrate the protoplast (i.e. the interior of the cell), but invaginate the cell membrane. The structure of the arbuscules greatly increases the contact surface area between the hypha and the cell cytoplasm to facilitate the transfer of nutrients between them.

Arbuscular mycorrhizas are formed only by fungi in the division Glomeromycota. Fossil evidence[23] and DNA sequence analysis[28] suggest that this mutualism appeared 400-460 million years ago, when the first plants were colonizing land. Arbuscular mycorrhizas are found in 85% of all plant families, and occur in many crop species.[24] The hyphae of arbuscular mycorrhizal fungi produce the glycoprotein glomalin, which may be one of the major stores of carbon in the soil. Arbuscular mycorrhizal fungi have (possibly) been asexual for many millions of years and, unusually, individuals can contain many genetically different nuclei (a phenomenon called heterokaryosis).[29]

Ectomycorrhiza[edit]

Ectomycorrhizal beech
Main article: Ectomycorrhiza

Ectomycorrhizas, or EcM, are typically formed between the roots of around 10% of plant families, mostly woody plants including the birch, dipterocarp, eucalyptus, oak, pine, and rose[24] families, orchids,[30] and fungi belonging to the Basidiomycota, Ascomycota, and Zygomycota. Some EcM fungi, such as many Leccinum and Suillus, are symbiotic with only one particular genus of plant, while other fungi, such as the Amanita, are generalists that form mycorrhizas with many different plants.[31] An individual tree may have 15 or more different fungal EcM partners at one time.[32] Thousands of ectomycorrhizal fungal species exist, hosted in over 200 genera. A recent study has conservatively estimated global ectomycorrhizal fungal species richness at approximately 7750 species. Although, on the basis of estimates of knowns and unknowns in macromycete diversity, a final estimate of ECM species richness would likely be between 20000 and 25000.[33]

Ectomycorrhizas consist of a hyphal sheath, or mantle, covering the root tip and a Hartig net of hyphae surrounding the plant cells within the root cortex. In some cases the hyphae may also penetrate the plant cells, in which case the mycorrhiza is called an ectendomycorrhiza. Outside the root, Ectomycorrhizal extramatrical mycelium forms an extensive network within the soil and leaf litter.

Nutrients can be shown to move between different plants through the fungal network. Carbon has been shown to move from paper birch trees into Douglas-fir trees thereby promoting succession in ecosystems.[34] The ectomycorrhizal fungus Laccaria bicolor has been found to lure and kill springtails to obtain nitrogen, some of which may then be transferred to the mycorrhizal host plant. In a study by Klironomos and Hart, Eastern White Pine inoculated with L. bicolor was able to derive up to 25% of its nitrogen from springtails.[35][36]

The first genomic sequence for a representative of symbiotic fungi, the ectomycorrhizal basidiomycete Laccaria bicolor, has been published.[37] An expansion of several multigene families occurred in this fungus, suggesting that adaptation to symbiosis proceeded by gene duplication. Within lineage-specific genes those coding for symbiosis-regulated secreted proteins showed an up-regulated expression in ectomycorrhizal root tips suggesting a role in the partner communication. Laccaria bicolor is lacking enzymes involved in the degradation of plant cell wall components (cellulose, hemicellulose, pectins and pectates), preventing the symbiont from degrading host cells during the root colonisation. By contrast, Laccaria bicolor possesses expanded multigene families associated with hydrolysis of bacterial and microfauna polysaccharides and proteins. This genome analysis revealed the dual saprotrophic and biotrophic lifestyle of the mycorrhizal fungus that enables it to grow within both soil and living plant roots.

Ericoid mycorrhiza[edit]

An ericoid mycorrhizal fungus isolated from Woollsia pungens[38]
Main article: Ericoid mycorrhiza

Ericoid mycorrhizas are the third of the three more ecologically important types. They have a simple intraradical (grow in cells) phase, consisting of dense coils of hyphae in the outermost layer of root cells. There is no periradical phase and the extraradical phase consists of sparse hyphae that don't extend very far into the surrounding soil. They might form sporocarps (probably in the form of small cups), but their reproductive biology is little understood.[25]

Ericoid mycorrhizas have also been shown to have considerable saprotrophic capabilities, which would enable plants to receive nutrients from not-yet-decomposed materials via the decomposing actions of their ericoid partners.[39]

Discovery[edit]

Associations of fungi with the roots of plants have been known since at least the mid-19th century. However early observers simply recorded the fact without investigating the relationships between the two organisms.[40] This symbiosis was studied and described by Franciszek Kamieński in 1879–1882.[41] Further research was carried out by Albert Bernhard Frank, who introduced the term mycorrhiza in 1885.[42]

See also[edit]

References[edit]

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External links[edit]