Mycorrhiza: Difference between revisions

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Introductory video animation providing some basic information about mycorrhizas.

A mycorrhiza (Template:Etymology; Template:Plural form, mycorrhiza, or mycorrhizas)^[1] is a symbiotic association between a fungus and a plant,^[2] in which fungal hyphae and plant roots become interconnected and form an interface on the cellular level.^[3]^[4]^[5] The term mycorrhiza refers to the role of the fungus in the plant's rhizosphere, the plant root system and its surroundings. Mycorrhizae play important roles in plant nutrition, soil biology, and soil chemistry.

In a mycorrhizal association, the fungus colonizes the host plant's root tissues, either intracellularly as in arbuscular mycorrhizal fungi, or extracellularly as in ectomycorrhizal fungi.^[6] The association is normally mutualistic. In particular species, or in particular circumstances, mycorrhizae may have a parasitic association with host plants.^[7]

Definition

A mycorrhiza is a symbiotic association between a green plant and a fungus. The plant makes organic molecules by photosynthesis and supplies them to the fungus in the form of sugars or lipids, while the fungus supplies the plant with water and mineral nutrients, such as phosphorus, nitrogen, or zinc,^[8] taken from the soil. Mycorrhizas are located in the roots of vascular plants, but mycorrhiza-like associations also occur in bryophytes^[9] and there is fossil evidence that early land plants that lacked roots formed arbuscular mycorrhizal associations.^[10] Most plant species form mycorrhizal associations, though some families like Brassicaceae and Chenopodiaceae cannot. Different forms for the association are detailed in the next section. The most common is the arbuscular type that is present in 70% of plant species, including many crop plants such as cereals and legumes.^[11]

Evolution

Emergence alongside terrestrial plants

Fossil and genetic evidence indicate that mycorrhizae emerged as early as 450-500 million years ago, potentially between fungus-like protists and algae. Arbuscular mycorrhizal relationships appeared earliest, coinciding with the terrestrialization of plants.^[12] Genetic evidence indicates that all land plants share a single common ancestor,^[13] which appears to have quickly adopted mycorrhizal symbiosis, and research suggests that proto-mycorrhizal fungi were a key factor enabling plant terrestrialization.^[14] There is a strong consensus among paleomycologists that mycorrhizal fungi served as a primitive root system for early terrestrial plants.^[12] This is because, prior to plant colonization of land, soils were nutrient sparse and plants had yet to develop root systems. Without complex root systems, early terrestrial plants would have been incapable of absorbing recalcitrant ions from mineral substrates, such as phosphate, a key nutrient for plant growth.^[15]

Fossil record and genomic analysis

Fossils of Glomeromycotan spores and hyphae date to 460 million years ago, but the fossils were not associated with plants. The earliest terrestrial communities were similar to modern biocrusts. Lichen-like associations between fungi and cyanobacteria were an important part of these communities. The first land plants were similar to mosses, with simple vascular systems and no leaves or roots.^[16]

The earliest direct fossil evidence of early mycorrhizal symbiosis is found in the 407 million year old Rhynie chert, which contains an assemblage of "exceptionally preserved" fossil plants colonized by multiple para-mycorrhizal fungi.^[10] The Rhynie chert fossils show Glomeromycotan and Mucoromycotan fungi engaged in mycorrhiza-like associations with cells of the plants Aglaophyton major and Horneophyton lignieri. The fossils suggest a mutualistic association between the plants and the colonizing fungi, because distinctive nutrient exchange structures (arbuscules and hyphal coils, in Glomeromycotina and Mucoromycotina respectively) are preserved and the colonized cells appear to have been alive at the time of infection by the fungus.^[16]

These early associations are referred to as mycorrhiza-like or para-mycorrhizal because mycorrhiza are defined by the fungal colonization of plant roots, and early plants did not have any roots. The earliest fossils of arbuscular mycorrhizal fungi in plant roots originate from 315-303 million years ago, and show fungi belonging to Glomeromycotina in the root systems of a giant lycophyte, Lepidodendron, and an early relative of the conifers, Cordaites.^[16] Arbuscular mycorrhizal fungi were found in 240 million year old fossils of Antarcticycas schopfii.

Ectomycorrhizae developed substantially later, during the Jurassic period, while most other modern forms of mycorrhizal symbiosis, including orchid and ericoid mycorrhizae, date to the period of angiosperm radiation in the Cretaceous period.^[17] Ectomycorrhizae appear in the fossil record 48.7 million years ago, in the Eocene, with a fossil of ectomycorrhizal fungi colonizing Pinus roots.^[18] However, it is believed that the first ectomycorrhizal relationships evolved in the stem group Pinaceae around the radiation of the Pinaceae crown group in the mid Jurassic, 175 million or so years ago.^[18]

Fossils preservation of Ericoid mycorrhizae and orchid mycorrhizae is lacking. Calibrated molecular phylogeny is used to estimate when these mycorrhizal types originated.^[18] The origins of orchid mycorrhizae are unclear, though orchids themselves are thought to have originated in the Cretaceous period. Ericoid mycorrhizae are estimated to have the most recent evolutionary origins of mycorrhizal types, evolving around 118 million years ago from free-living saprotrophic ancestors.^[19] Ericoid mycorrhizal fungi evolved from multiple lineages of fungi, primarily ascomycetes from the Leotiomycetes, as well as basidiomycetes from the family Serendipitaceae.^[20]

Origins in plants

In plants, the genes for forming mycorrhizal symbiosis are highly conserved and originate from a common ancestor, meaning that the ability to form mycorrhizae is ancestral to all land plants.^[21] Non-mycorrhizal plant lineages, such as the Brassicaceae, lost the ability to form mycorrhizae at some point in their evolution.^[22] The earliest mycorrhizae were arbuscular mycorrhizae, and other forms, such as ectomycorrhizae and orchid mycorrhizae, evolved when plant hosts switched from symbiosis with Glomeromycotina to symbiosis with different fungal lineages.^[23]

There is genetic evidence that the symbiosis between legumes and nitrogen-fixing bacteria is derived from mycorrhizal symbiosis.^[24] The modern distribution of mycorrhizal fungi appears to reflect an increasing complexity and competition in root morphology associated with the dominance of angiosperms in the Cenozoic Era, characterized by complex ecological dynamics between species.^[25]

Origins in fungi

In fungi, mycorrhizal symbiosis had multiple independent origins among different lineages of fungi. Arbuscular mycorrhizal fungi form their own monophyletic phylum, whereas other mycorrhizal fungi convergently evolved similar lifestyles.

Arbuscular mycorrhizae

The phylum Glomeromycota, which forms the arbuscular mycorrhizal symbiosis, is the oldest mycorrhizal lineage. The arbuscular mycorrhizal symbiosis evolved only once in fungi; all arbuscular mycorrhizal fungi belong to Glomeromycota and share a common ancestor.^[19] 244 species have been identified based on differences in the appearance of their spores, but genetic studies suggest that 300-1600 species may exist in Glomeromycota.^[21] All members of Glomeromycota are obligate biotrophs, entirely dependent upon their plant hosts for survival.^[26] Arbuscular mycorrhizal fungi are considered to be generalists, with minimal host plant specificity. AM symbiosis has been observed in almost every seed plant taxonomic division, or around 67% of species.^[15] Arbuscular mycorrhizae take on most angiosperms, some gymnosperms, pteridophytes, and nonvascular plants as plant hosts.^[27] Arbuscular mycorrhizae have been observed in the seedling stage of otherwise ectomycorrhizal partners, suggesting that arbuscular mycorrhizal fungi may be able to infect almost any land plant given proper circumstances.^[28]

Other forms of mycorrhizal symbiosis, such as ectomycorrhizae, orchid mycorrhizae, and ericoid mycorrhizae, emerged multiple times in different lineages of fungi through convergent evolution. Unlike arbuscular mycorrhizal fungi, some of these fungi are only facultatively symbiotic, and can live by themselves without a plant host under some conditions.

Ectomycorrhizae

Ectomycorrhizal fungi evolved from free-living saprotrophs, mostly in Basidiomycota and Ascomycota, and some became dependent on plant hosts when they lost genes necessary for decaying lignin and other plant materials.^[21] There are 20,000 to 25,000 species of ectomycorrhizal fungi, but only 6,000 to 7,000 plant species that form ectomycorrhizal symbiosis.^[29] In angiosperms, it is believed that ectomycorrhizal partnerships have developed independently at least 18 times, and in fungi, around 80 times.^[18]^[19] The main evolutionary driver for ectomycorrhizae is switching of nutritional modes from saprotrophs.^[30] Phylogenomic analysis of various ectomycorrhizal fungal genomes has confirmed the convergent evolution of ectomycorrhizal fungi from white and brown-rot fungi, as well as from soil saprotrophs.^[30]^[28] Some lineages of ectomycorrhizae have likely evolved from endophytic ancestors, fungi that live within plants without damaging them.^[30] Some ectomycorrhizal fungi have gone through apparent evolutionary reversal back into saprotrophic ecology. This is possible because some lineages of ectomycorrhizal fungi retain enzymes for breaking down lignin.^[31]

Orchid mycorrhizae

Orchid mycorrhizal fungi, which mostly originate from Ascomycota and Basidiomycota, are less understood. Some fungi that participate in orchid mycorrhizal symbiosis can also form ectomycorrhizal symbiosis with other plants, or live independently of a plant host. Some orchid mycorrhizal fungi can also live as plant pathogens.^[23]^[21]

Ericoid mycorrhizae

Ericoid mycorrhizal associations have the most recent origins and the lowest species richness among both plant and fungal partners.^[19] This specialization suggests that ericoid mycorrhizal partners evolved in parallel with one another in response to environmental change, rather than through reciprocal species-to-species level selection.^[32] Ericoid mycorrhizal relationships are found in extremely nutrient poor soils in the northern and southern hemispheres.^[31] These environments of low mineral nutrient availability have led to native plants developing sclerophylly, where plants become high in lignin and low in phosphorus and nitrogen.^[31] As a result, decaying plant matter in these areas has an abnormally high carbon to nitrogen ratio, making it resistant to microbial decay. Ericoid mycorrhizae have apparently evolved to conserve minerals in nutrient deficient sclerophyllous litter by directly cycling these nutrients throughout the mycorrhiza system.^[31] Ericoid mycorrhizae also retain saprotrophic abilities, allowing them to extract nitrogen and phosphorus from unmineralized organic material, and resist negative outcomes from high concentrations of toxic cations in the acidic soil environment.^[31]

Types

The mycorrhizal lifestyle has independently convergently evolved multiple times in the history of Earth.^[33] There are multiple ways to categorize mycorrhizal symbiosis. The largest division is between 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.^[34]^[35]

Similar symbiotic relationships

Some forms of plant-fungal symbiosis are similar to mycorrhizae, but considered distinct. One example is fungal endophytes. Endophytes are defined as organisms that can live within plant cells without causing harm to the plant. They are distinguishable from mycorrhizal fungi by the absence of nutrient-transferring structures for bringing in nutrients from outside the plant.^[33] Some lineages of mycorrhizal fungi may have evolved from endophytes into mycorrhizal fungi,^[36] and some fungi can live as mycorrhizae or as endophytes.

Ectomycorrhiza

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Beech is ectomycorrhizal

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Leccinum aurantiacum, an ectomycorrhizal fungus

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Ectomycorrhizae are distinct in that they do not penetrate into plant cells, but instead form a structure called a Hartig net that penetrates between cells.^[37] Ectomycorrhizas consist of a hyphal sheath, or mantle, covering the root tip and the 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 endomycorrhiza. Outside the root, ectomycorrhizal extramatrical mycelium forms an extensive network within the soil and leaf litter. Other forms of mycorrhizae, including arbuscular, ericoid, arbutoid, monotropoid, and orchid mycorrhizas, are considered endomycorrhizae.^[38]

Ectomycorrhizas, or EcM, are symbiotic associations between the roots of around 10% of plant families, mostly woody plants including the birch, dipterocarp, eucalyptus, oak, pine, and rose^[39] families, orchids,^[40] and fungi belonging to the Basidiomycota, Ascomycota, and Zygomycota. Ectomycorrhizae associate with relatively few plant species, only about 2% of plant species on Earth, but the species they associate with are mostly trees and woody plants that are highly dominant in their ecosystems, meaning plants in ectomycorrhizal relationships make up a large proportion of plant biomass.^[41] 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.^[42] An individual tree may have 15 or more different fungal EcM partners at one time.^[43] While the diversity of plants involved in EcM is low, the diversity of fungi involved in EcM is high. 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 probably be between 20,000 and 25,000.^[44] Ectomycorrhizal fungi evolved independently from saprotrophic ancestors many times in the group's history.^[45]

Nutrients can be shown to move between different plants through the fungal network. Carbon has been shown to move from paper birch seedlings into adjacent Douglas-fir seedlings, although not conclusively through a common mycorrhizal network,^[46] thereby promoting succession in ecosystems.^[47] 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.^[48]^[49] When compared with non-mycorrhizal fine roots, ectomycorrhizae may contain very high concentrations of trace elements, including toxic metals (cadmium, silver) or chlorine.^[50]

The first genomic sequence for a representative of symbiotic fungi, the ectomycorrhizal basidiomycete L. bicolor, was published in 2008.^[51] 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. L. 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, L. 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. Since then, the genomes of many other ectomycorrhizal fungal species have been sequenced further expanding the study of gene families and evolution in these organisms.^[52]

Arbutoid mycorrhiza

This type of mycorrhiza involves plants of the Ericaceae subfamily Arbutoideae. It is however different from ericoid mycorrhiza and resembles ectomycorrhiza, both functionally and in terms of the fungi involved.^[53] It differs from ectomycorrhiza in that some hyphae actually penetrate into the root cells, making this type of mycorrhiza an ectendomycorrhiza.^[54]

Arbuscular mycorrhiza

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Wheat has arbuscular mycorrhiza.

Arbuscular mycorrhizas, (formerly known as vesicular-arbuscular mycorrhizas), have hyphae that penetrate plant cells, producing branching, tree-like structures called arbuscules within the plant cells for nutrient exchange. Often, balloon-like storage structures, termed vesicles, are also produced. In this interaction, fungal hyphae do not in fact penetrate the protoplast (i.e. the interior of the cell), but invaginate the cell membrane, creating a so-called peri-arbuscular membrane. The structure of the arbuscules greatly increases the contact surface area between the hypha and the host cell cytoplasm to facilitate the transfer of nutrients between them. Arbuscular mycorrhizas are obligate biotrophs, meaning that they depend upon the plant host for both growth and reproduction; they have lost the ability to sustain themselves by decomposing dead plant material.^[55] Twenty percent of the photosynthetic products made by the plant host are consumed by the fungi, the transfer of carbon from the terrestrial host plant is then exchanged by equal amounts of phosphate from the fungi to the plant host.^[56]

Contrasting with the pattern seen in ectomycorrhizae, the species diversity of AMFs is very low, but the diversity of plant hosts is very high; an estimated 78% of all plant species associate with AMFs.^[41] Arbuscular mycorrhizas are formed only by fungi in the division Glomeromycota. Fossil evidence^[10] and DNA sequence analysis^[57] 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.^[39] The hyphae of arbuscular mycorrhizal fungi produce the glycoprotein glomalin, which may be one of the major stores of carbon in the soil.^[58] 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).^[59]

Mucoromycotina fine root endophytes

Mycorrhizal fungi belonging to Mucoromycotina, known as "fine root endophytes" (MFREs), were mistakenly identified as arbuscular mycorrhizal fungi until recently. While similar to AMF, MFREs are from subphylum Mucoromycotina instead of Glomeromycotina. Their morphology when colonizing a plant root is very similar to AMF, but they form fine textured hyphae.^[37] Effects of MFREs may have been mistakenly attributed to AMFs due to confusion between the two, complicated by the fact that AMFs and MFREs often colonize the same hosts simultaneously. Unlike AMFs, they appear capable of surviving without a host. This group of mycorrhizal fungi is little understood, but appears to prefer wet, acidic soils and forms symbiotic relationships with liverworts, hornworts, lycophytes, and angiosperms.^[60]

Ericoid mycorrhiza

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An ericoid mycorrhizal fungus isolated from Woollsia pungens^[61]

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Ericoid mycorrhizae, or ErMs, involve only plants in Ericales and are the most recently evolved of the major mycorrhizal relationships. Plants that form ericoid mycorrhizae are mostly woody understory shrubs; hosts include blueberries, bilberries, cranberries, mountain laurels, rhododendrons, heather, neinei, and giant grass tree. ErMs are most common in boreal forests, but are found in two-thirds of all forests on Earth.^[41] Ericoid mycorrhizal fungi belong to several different lineages of fungi. Some species can live as endophytes entirely within plant cells even within plants outside the Ericales, or live independently as saprotrophs that decompose dead organic matter. This ability to switch between multiple lifestyle types makes ericoid mycorrhizal fungi very adaptable.^[33]

Plants that participate in these symbioses have specialized roots with no root hairs, which are covered with a layer of epidermal cells that the fungus penetrates into and completely occupies.^[37] The fungi have a simple intraradical (growth 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 poorly understood.^[35]

Plants participating in ericoid mycorrhizal symbioses are found in acidic, nutrient-poor conditions.^[33] Whereas AMFs have lost their saprotrophic capabilities, and EcM fungi have significant variation in their ability to produce enzymes needed for a saprotrophic lifestyle,^[41] fungi involved in ErMs have fully retained the ability to decompose plant material for sustenance. Some ericoid mycorrhizal fungi have actually expanded their repertoire of enzymes for breaking down organic matter. They can extract nitrogen from cellulose, hemicellulose, lignin, pectin, and chitin. This would increase the benefit they can provide to their plant symbiotic partners.^[62]

Orchid mycorrhiza

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All orchids are myco-heterotrophic at some stage during their lifecycle, meaning that they can survive only if they form orchid mycorrhizae. Orchid seeds are so small that they contain no nutrition to sustain the germinating seedling, and instead must gain the energy to grow from their fungal symbiont.^[37] The OM relationship is asymmetric; the plant seems to benefit more than the fungus, and some orchids are entirely mycoheterotrophic, lacking chlorophyll for photosynthesis. It is actually unknown whether fully autotrophic orchids that do not receive some of their carbon from fungi exist or not.^[63] Like fungi that form ErMs, OM fungi can sometimes live as endophytes or as independent saprotrophs. In the OM symbiosis, hyphae penetrate into the root cells and form pelotons (coils) for nutrient exchange.

Monotropoid mycorrhiza

'Monotropa' plant unable to photosynthesis, collects food from monotropoid mycorrhiza

Monotropa plant unable to photosynthesis, collects food from monotropoid mycorrhiza

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This type of mycorrhiza occurs in the subfamily Monotropoideae of the Ericaceae, as well as several genera in the Orchidaceae. These plants are heterotrophic or mixotrophic and derive their carbon from the fungus partner. This is thus a non-mutualistic, parasitic type of mycorrhizal symbiosis.Script error: No such module "Unsubst".

Function

File:Mycorrhizal network.svg

Nutrient exchanges and communication between a mycorrhizal fungus and plants.

Mycorrhizal fungi form a mutualistic relationship with the roots of most plant species. In such a relationship, both the plants themselves and those parts of the roots that host the fungi, are said to be mycorrhizal. Relatively few of the mycorrhizal relationships between plant species and fungi have been examined to date, but 95% of the plant families investigated are predominantly mycorrhizal either in the sense that most of their species associate beneficially with mycorrhizae, or are absolutely dependent on mycorrhizae. The Orchidaceae are notorious as a family in which the absence of the correct mycorrhizae is fatal even to germinating seeds.^[64]

Recent research into ectomycorrhizal plants in boreal forests has indicated that mycorrhizal fungi and plants have a relationship that may be more complex than simply mutualistic. This relationship was noted when mycorrhizal fungi were unexpectedly found to be 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. They go on to explain how this updated model could explain 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.^[65] It has also been suggested that evolutionary and phylogenetic relationships can explain much more variation in the strength of mycorrhizal mutualisms than ecological factors.^[66]

File:Mutualistic mycorrhiza en.svg

Within mycorrhiza, the plant gives carbohydrates (products of photosynthesis) to the fungus, while the fungus gives the plant water and minerals.

Formation

To successfully engage in mutualistic symbiotic relationships with other organisms, such as mycorrhizal fungi and any of the thousands of microbes that colonize plants, plants must discriminate between mutualists and pathogens, allowing the mutualists to colonize while activating an immune response towards the pathogens. Plant genomes code for potentially hundreds of receptors for detecting chemical signals from other organisms. Plants dynamically adjust their symbiotic and immune responses, changing their interactions with their symbionts in response to feedbacks detected by the plant.^[67] In plants, the mycorrhizal symbiosis is regulated by the common symbiosis signaling pathway (CSSP), a set of genes involved in initiating and maintaining colonization by endosymbiotic fungi and other endosymbionts such as Rhizobia in legumes. The CSSP has origins predating the colonization of land by plants, demonstrating that the co-evolution of plants and arbuscular mycorrhizal fungi is over 500 million years old.^[45] In arbuscular mycorrhizal fungi, the presence of strigolactones, a plant hormone, secreted from roots induces fungal spores in the soil to germinate, stimulates their metabolism, growth and branching, and prompts the fungi to release chemical signals the plant can detect.^[68] Once the plant and fungus recognize one another as suitable symbionts, the plant activates the common symbiotic signaling pathway, which causes changes in the root tissues that enable the fungus to colonize.^[69]

Experiments with arbuscular mycorrhizal fungi have identified numerous chemical compounds to be involved in the "chemical dialog" that occurs between the prospective symbionts before symbiosis is begun. In plants, almost all plant hormones play a role in initiating or regulating AMF symbiosis, and other chemical compounds are also suspected to have a signaling function. While the signals emitted by the fungi are less understood, it has been shown that chitinaceous molecules known as Myc factors are essential for the formation of arbuscular mycorrhizae. Signals from plants are detected by LysM-containing receptor-like kinases, or LysM-RLKs. AMF genomes also code for potentially hundreds of effector proteins, of which only a few have a proven effect on mycorrhizal symbiosis, but many others likely have a function in communication with plant hosts as well.^[68]

Many factors are involved in the initiation of mycorrhizal symbiosis, but particularly influential is the plant's need for phosphorus. Experiments involving rice plants with a mutation disabling their ability to detect P starvation show that arbuscular mycorrhizal fungi detection, recruitment and colonization is prompted when the plant detects that it is starved of phosphorus.^[60] Nitrogen starvation also plays a role in initiating AMF symbiosis.^[68]

Mechanisms

The mechanisms by which mycorrhizae increase absorption include some that are physical and some that are chemical. Physically, most mycorrhizal mycelia are much smaller in diameter than the smallest root or root hair, and thus can explore soil material that roots and root hairs cannot reach, and provide a larger surface area for absorption. Chemically, the cell membrane chemistry of fungi differs from that of plants. For example, they may secrete organic acids that dissolve or chelate many ions, or release them from minerals by ion exchange.^[70] Mycorrhizae are especially beneficial for the plant partner in nutrient-poor soils.^[71]

Sugar-water/mineral exchange

File:Mycorrhiza.svg

In this mutualism, fungal hyphae (E) increase the surface area of the root and uptake of key nutrients while the plant supplies the fungi with fixed carbon (A=root cortex, B=root epidermis, C=arbuscle, D=vesicle, F=root hair, G=nuclei).

The mycorrhizal mutualistic association provides the fungus with relatively constant and direct access to carbohydrates, such as glucose and sucrose.^[72] 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, partly because of the large surface area of fungal hyphae, which are much longer and finer than plant root hairs, and partly because some such fungi can mobilize soil minerals unavailable to the plants' roots. The effect is thus to improve the plant's mineral absorption capabilities.^[73]

Unaided plant roots may be unable to take up nutrients that are chemically or physically immobilised; examples include phosphate ions and micronutrients such as iron. One form of such immobilization occurs in soil with high clay content, or soils with a strongly basic pH. The mycelium of the mycorrhizal fungus can, however, access many such nutrient sources, and make them available to the plants they colonize.^[74] Thus, many plants are able to obtain phosphate without using soil as a source. Another form of immobilisation is when nutrients are locked up in organic matter that is slow to decay, such as wood, and some mycorrhizal fungi act directly as decay organisms, mobilising the nutrients and passing some onto the host plants; for example, in some dystrophic forests, large amounts of phosphate and other nutrients are taken up by mycorrhizal hyphae acting directly on leaf litter, bypassing the need for soil uptake.^[75] Inga alley cropping, an agroforestry technique proposed as an alternative to slash and burn rainforest destruction,^[76] relies upon mycorrhiza within the root system of species of Inga to prevent the rain from washing phosphorus out of the soil.^[77]

In some more complex relationships, mycorrhizal fungi do not just collect immobilised soil nutrients, but connect individual plants together by mycorrhizal networks that transport water, carbon, and other nutrients directly from plant to plant through underground hyphal networks.^[78]

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.^[79]

Disease, drought and salinity resistance and its correlation to mycorrhizae

Mycorrhizal plants are often more resistant to diseases, such as those caused by microbial soil-borne pathogens. These associations have been found to assist in plant defense both above and belowground. Mycorrhizas have been found to excrete enzymes that are toxic to soil borne organisms such as nematodes.^[80] More recent studies have shown that mycorrhizal associations result in a priming effect of plants that essentially acts as a primary immune response. When this association is formed a defense response is activated similarly to the response that occurs when the plant is under attack. As a result of this inoculation, defense responses are stronger in plants with mycorrhizal associations.^[81] Ecosystem services provided by mycorrhizal fungi may depend on the soil microbiome.^[82] Furthermore, mycorrhizal fungi was significantly correlated with soil physical variable, but only with water level and not with aggregate stability^[83]^[84] and can lead also to more resistant to the effects of drought.^[85]^[86]^[87] Moreover, the significance of mycorrhizal fungi also includes alleviation of salt stress and its beneficial effects on plant growth and productivity. Although salinity can negatively affect mycorrhizal fungi, many reports show improved growth and performance of mycorrhizal plants under salt stress conditions.^[88]

Resistance to insects

Plants connected by mycorrhizal fungi in mycorrhizal networks can use these underground connections to communicate warning signals.^[89]^[90] For example, when a host plant is attacked by an aphid, the plant signals surrounding connected plants of its condition. Both the host plant and those connected to it release volatile organic compounds that repel aphids and attract parasitoid wasps, predators of aphids.^[89] This assists the mycorrhizal fungi by conserving its food supply.^[89]

Colonization of barren soil

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.^[91] The absence of mycorrhizal fungi can also slow plant growth in early succession or on degraded landscapes.^[92] The introduction of alien mycorrhizal plants to nutrient-deficient ecosystems puts indigenous non-mycorrhizal plants at a competitive disadvantage.^[93] This aptitude to colonize barren soil is defined by the category Oligotroph.

Resistance to toxicity

Fungi 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.^[94] 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.^[93]

Occurrence of mycorrhizal associations

Mycorrhizas are present in 92% of plant families studied (80% of species),^[39] with arbuscular mycorrhizas being the ancestral and predominant form,^[39] and the most prevalent symbiotic association found in the plant kingdom.^[72] The structure of arbuscular mycorrhizas has been highly conserved since their first appearance in the fossil record,^[10] with both the development of ectomycorrhizas and the loss of mycorrhizas, evolving convergently on multiple occasions.^[39]

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.^[95] This symbiosis was studied and described by Franciszek Kamieński in 1879–1882.^[96]^[97]

Climate change

CO₂ released by human activities is causing climate change and possible damage to mycorrhizae, but the direct effect of an increase in the gas should be to benefit plants and mycorrhizae.^[98] In Arctic regions, nitrogen and water are harder for plants to obtain, making mycorrhizae crucial to plant growth.^[99] Since mycorrhizae tend to do better in cooler temperatures, warming could be detrimental to them.^[100] Gases such as SO₂, NO_x, and O₃ produced by human activity may harm mycorrhizae, causing reduction in "propagules, the colonization of roots, degradation in connections between trees, reduction in the mycorrhizal incidence in trees, and reduction in the enzyme activity of ectomycorrhizal roots."^[101]

A company in Israel, Groundwork BioAg, has discovered a method of using mycorrhizal fungi to increase agricultural crops while sequestering greenhouse gases and eliminating CO₂ from the atmosphere.^[102]

Conservation and mapping

In 2021, the Society for the Protection of Underground Networks was launched. SPUN is a science-based initiative to map and protect the mycorrhizal networks regulating Earth's climate and ecosystems. Its stated goals are mapping, protecting, and harnessing mycorrhizal fungi.

References

Template:Reflist

External links

Template:Americana Poster

International Mycorrhiza Society International Mycorrhiza Society
Mohamed Hijri: A simple solution to the coming phosphorus crisis video recommending agricultural mycorrhiza use to conserve phosphorus reserves & 85% waste problem @Ted.com
Mycorrhizal Associations: The Web Resource Comprehensive illustrations and lists of mycorrhizal and nonmycorrhizal plants and fungi
Mycorrhizas – a successful symbiosis Biosafety research into genetically modified barley
MycorWiki a portal concerned with the biology and ecology of ectomycorrhizal fungi and other forest fungi.

Template:Authority control

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↑ Harley, J. L.; Smith, S. E. 1983. Mycorrhizal symbiosis (1st ed.). Academic Press, London.
↑ ^a ^b Allen, Michael F. 1991. The ecology of mycorrhizae. Cambridge University Press, Cambridge.
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↑ he Israeli Company That Uses Fungus to Tackle the Climate and Soil Crises, Haaretz

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[34] Harley, J. L.; Smith, S. E. 1983. Mycorrhizal symbiosis (1st ed.). Academic Press, London.

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[102] Israeli Company That Uses Fungus to Tackle the Climate and Soil Crises, Haaretz

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@@ Line 8: / Line 8: @@
 [[File:Mycorrhiza I – Mycorrhiza and the Environment.webm|thumb|Introductory video animation providing some basic information about mycorrhizas.]]
-A '''mycorrhiza''' ({{etymology|grc|''{{Wikt-lang|grc|μύκης}}'' ({{grc-transl|μύκης}})|fungus||''{{Wikt-lang|grc|ῥίζα}}'' ({{grc-transl|ῥίζα}})|root}}; {{plural form|'''mycorrhizae'''}}, '''mycorrhiza''', or '''mycorrhizas''')<ref>{{cite web |last1=Deacon |first1=Jim |title=The Microbial World: Mycorrhizas |url=http://archive.bio.ed.ac.uk/jdeacon/microbes/mycorrh.htm |url-status=dead |archive-url=https://web.archive.org/web/20180427082137/http://archive.bio.ed.ac.uk/jdeacon/microbes/mycorrh.htm |archive-date=2018-04-27 |access-date=11 January 2019 |website=bio.ed.ac.uk (archived)}}</ref> is a [[symbiosis|symbiotic]] association between a [[fungus]] and a [[plant]].<ref>{{Cite book |last1=Kirk |first1=P. M. |first2=P. F. |last2=Cannon |first3=J. C. |last3=David |first4=J. |last4=Stalpers |date=2001 |title=Ainsworth and Bisby's Dictionary of the Fungi |edition=9th |publisher=CAB International |location=Wallingford, UK }}</ref> The term mycorrhiza refers to the role of the fungus in the plant's [[rhizosphere]], the plant [[root]] system and its surroundings. Mycorrhizae play important roles in [[plant nutrition]], [[soil life|soil biology]], and [[soil chemistry]].
+A '''mycorrhiza''' ({{etymology|grc|''{{Wikt-lang|grc|μύκης}}'' ({{grc-transl|μύκης}})|fungus||''{{Wikt-lang|grc|ῥίζα}}'' ({{grc-transl|ῥίζα}})|root}}; {{plural form|'''mycorrhizae'''}}, '''mycorrhiza''', or '''mycorrhizas''')<ref>{{cite web |last1=Deacon |first1=Jim |title=The Microbial World: Mycorrhizas |url=http://archive.bio.ed.ac.uk/jdeacon/microbes/mycorrh.htm |archive-url=https://web.archive.org/web/20180427082137/http://archive.bio.ed.ac.uk/jdeacon/microbes/mycorrh.htm |archive-date=2018-04-27 |access-date=11 January 2019 |website=bio.ed.ac.uk (archived)}}</ref> is a [[symbiosis|symbiotic]] association between a [[fungus]] and a [[plant]],<ref>{{Cite book |last1=Kirk |first1=P. M. |first2=P. F. |last2=Cannon |first3=J. C. |last3=David |first4=J. |last4=Stalpers |date=2001 |title=Ainsworth and Bisby's Dictionary of the Fungi |edition=9th |publisher=CAB International |location=Wallingford, UK }}</ref> in which fungal hyphae and plant roots become interconnected and form an interface on the cellular level.<ref>{{cite book |last1=Read |first1=D.J. |editor1-last=Varma |editor1-first=Ajit |editor2-last=Hock |editor2-first=Bertold |title=Mycorrhiza: Structure, Function, Molecular Biology and Biotechnology |date=2013 |publisher=Springer Science & Business Media |isbn=978-3662037799 |pages=3–5 |chapter=Mycorrhiza- the state of the art}}</ref><ref>{{cite journal |last1=Brundrett |first1=Marc C |title=Diversity and classification of mycorrhizal associations |journal=Biological Reviews of the Cambridge Philosophical Society |date=2004 |volume=79 |issue=3 |pages=473–95 |doi=10.1017/S1464793103006316 |quote=“A new definition of mycorrhizas that encompasses all types of these associations while excluding other plant-fungus interactions is provided. This definition recognises the importance of nutrient transfer at an interface resulting from synchronised plant-fungus development.”}}</ref><ref>{{cite journal |last1=Balestrini |first1=Rafaella |last2=Bonfante |first2=Paola |title=Cell wall remodeling in mycorrhizal symbiosis: a way towards biotrophism |journal=Frontiers in Plant Science |date=2014 |volume=5 |page=237 |doi=10.3389/fpls.2014.00237 |bibcode=2014FrPS....5..237B |doi-access=free |quote=“All mycorrhizal interactions achieve full symbiotic functionality through the development of an extensive contact surface between the plant and fungal cells, where signals and nutrients are exchanged.”}}</ref> The term mycorrhiza refers to the role of the fungus in the plant's [[rhizosphere]], the plant [[root]] system and its surroundings. Mycorrhizae play important roles in [[plant nutrition]], [[soil life|soil biology]], and [[soil chemistry]].
-In a mycorrhizal association, the fungus colonizes the host plant's root tissues, either [[intracellular]]ly as in [[arbuscular mycorrhizal fungi]], or [[extracellular]]ly as in [[#Ectomycorrhiza|ectomycorrhizal]] fungi.<ref>{{Cite book |url=https://link.springer.com/book/10.1007/978-981-10-4115-0 |title=Arbuscular Mycorrhizas and Stress Tolerance of Plants |publisher=Springer Singapore |year=2017 |isbn=978-981-10-4115-0 |editor-last=Wu |editor-first=Qiang-Sheng |edition=1st |pages=1 |language=en |doi=10.1007/978-981-10-4115-0}}</ref> The association is normally [[Mutualism (biology)|mutualistic]]. In particular species, or in particular circumstances, mycorrhizae may have a [[Parasitism|parasitic]] association with host plants.<ref>{{cite journal |last1=Johnson |first1=N. C. |last2=Graham |first2=J. H. |last3=Smith |first3=F. A. |title=Functioning of mycorrhizal associations along the mutualism–parasitism continuum |journal=[[New Phytologist]] |date=1997 |doi=10.1046/j.1469-8137.1997.00729.x |volume=135 |issue=4 |pages=575–585|s2cid=42871574 |doi-access=free |bibcode=1997NewPh.135..575J }}</ref>
+In a mycorrhizal association, the fungus colonizes the host plant's root tissues, either [[intracellular]]ly as in [[arbuscular mycorrhizal fungi]], or [[extracellular]]ly as in [[#Ectomycorrhiza|ectomycorrhizal]] fungi.<ref>{{Cite book |url=https://link.springer.com/book/10.1007/978-981-10-4115-0 |title=Arbuscular Mycorrhizas and Stress Tolerance of Plants |publisher=Springer Singapore |year=2017 |isbn=978-981-10-4115-0 |editor-last=Wu |editor-first=Qiang-Sheng |edition=1st |page=1 |language=en |doi=10.1007/978-981-10-4115-0}}</ref> The association is normally [[Mutualism (biology)|mutualistic]]. In particular species, or in particular circumstances, mycorrhizae may have a [[Parasitism|parasitic]] association with host plants.<ref>{{cite journal |last1=Johnson |first1=N. C. |last2=Graham |first2=J. H. |last3=Smith |first3=F. A. |title=Functioning of mycorrhizal associations along the mutualism–parasitism continuum |journal=[[New Phytologist]] |date=1997 |doi=10.1046/j.1469-8137.1997.00729.x |volume=135 |issue=4 |pages=575–585|s2cid=42871574 |doi-access=free |bibcode=1997NewPh.135..575J }}</ref>
 == Definition ==
-A mycorrhiza is a symbiotic association between a green plant and a fungus. The plant makes organic molecules by [[photosynthesis]] and supplies them to the fungus in the form of sugars or lipids, while the fungus supplies the plant with water and mineral nutrients, such as [[phosphorus]], taken from the soil. Mycorrhizas are located in the roots of vascular plants, but mycorrhiza-like associations also occur in [[bryophytes]]<ref name=KottkeNebel>{{cite journal |date=2005 |first1=I. |last1=Kottke |first2=M. |last2=Nebel |title=The evolution of mycorrhiza-like associations in liverworts: An update |journal=New Phytologist |volume=167 |issue= 2 |pages=330–334 |doi=10.1111/j.1469-8137.2005.01471.x |pmid=15998388|doi-access=free |bibcode=2005NewPh.167..330K }}</ref> and there is fossil evidence that early land plants that lacked roots formed arbuscular mycorrhizal associations.<ref name="Remy et al."/> Most plant species form mycorrhizal associations, though some families like [[Brassicaceae]] and [[Chenopodiaceae]] cannot. Different forms for the association are detailed in the next section. The most common is the [[Arbuscular mycorrhiza|arbuscular type]] that is present in 70% of plant species, including many crop plants such as cereals and legumes.<ref>{{cite book |last1=Fortin |first1=J. André |display-authors=etal |title=Les Mycorhizes |date=2015 |publisher=Inra |location=Versaillles |isbn=978-2-7592-2433-3 |page=10 |edition=second}}</ref>
+A mycorrhiza is a symbiotic association between a green plant and a fungus. The plant makes organic molecules by [[photosynthesis]] and supplies them to the fungus in the form of sugars or lipids, while the fungus supplies the plant with water and mineral nutrients, such as [[phosphorus]], nitrogen, or zinc,<ref>{{Cite web |date=2017-04-01 |title=Mycorrhizal Fungi - Oklahoma State University |url=https://extension.okstate.edu/fact-sheets/mycorrhizal-fungi.html |access-date=2025-11-08 |website=extension.okstate.edu |language=en}}</ref> taken from the soil. Mycorrhizas are located in the roots of vascular plants, but mycorrhiza-like associations also occur in [[bryophytes]]<ref name=KottkeNebel>{{cite journal |date=2005 |first1=I. |last1=Kottke |first2=M. |last2=Nebel |title=The evolution of mycorrhiza-like associations in liverworts: An update |journal=New Phytologist |volume=167 |issue= 2 |pages=330–334 |doi=10.1111/j.1469-8137.2005.01471.x |pmid=15998388|doi-access=free |bibcode=2005NewPh.167..330K }}</ref> and there is fossil evidence that early land plants that lacked roots formed arbuscular mycorrhizal associations.<ref name="Remy et al."/> Most plant species form mycorrhizal associations, though some families like [[Brassicaceae]] and [[Chenopodiaceae]] cannot. Different forms for the association are detailed in the next section. The most common is the [[Arbuscular mycorrhiza|arbuscular type]] that is present in 70% of plant species, including many crop plants such as cereals and legumes.<ref>{{cite book |last1=Fortin |first1=J. André |display-authors=etal |title=Les Mycorhizes |date=2015 |publisher=Inra |location=Versaillles |isbn=978-2-7592-2433-3 |page=10 |edition=second}}</ref>
 == Evolution ==
-Fossil and genetic evidence indicate that mycorrhizae are ancient, potentially as old as the [[Timeline of plant evolution#Ordovician flora|terrestrialization of plants]]. Genetic evidence indicates that all land plants share a single common ancestor,<ref>{{Cite journal |last1=Harris |first1=Brogan J. |last2=Clark |first2=James W. |last3=Schrempf |first3=Dominik |last4=Szöllősi |first4=Gergely J. |last5=Donoghue |first5=Philip C. J. |last6=Hetherington |first6=Alistair M. |last7=Williams |first7=Tom A. |date=2022-09-29 |title=Divergent evolutionary trajectories of bryophytes and tracheophytes from a complex common ancestor of land plants |journal=Nature Ecology & Evolution |volume=6 |issue=11 |pages=1634–1643 |doi=10.1038/s41559-022-01885-x |pmc=9630106 |pmid=36175544|bibcode=2022NatEE...6.1634H }}</ref> which appears to have quickly adopted mycorrhizal symbiosis, and research suggests that proto-mycorrhizal fungi were a key factor enabling plant terrestrialization.<ref>{{Cite journal |last1=Puginier |first1=Camille |last2=Keller |first2=Jean |last3=Delaux |first3=Pierre-Marc |date=2022-08-29 |title=Plant–microbe interactions that have impacted plant terrestrializations |url=https://academic.oup.com/plphys/article/190/1/72/6596610 |journal=Plant Physiology |volume=190 |issue=1 |pages=72–84 |doi=10.1093/plphys/kiac258 |pmid=35642902 |pmc=9434271 }}</ref> The 400 million year old [[Rhynie chert]] contains an assemblage of fossil plants preserved in sufficient detail that arbuscular mycorrhizae have been observed in the stems of [[Aglaophyton|''Aglaophyton major'']], giving a lower bound for how late mycorrhizal symbiosis may have developed.<ref name="Remy et al." /> Ectomycorrhizae developed substantially later, during the [[Jurassic]] period, while most other modern mycorrhizal families, including orchid and ericoid mycorrhizae, date to the period of [[Flowering plant#Cretaceous|angiosperm radiation]] in the [[Cretaceous]] period.<ref>{{Cite journal |last1=Miyauchi |first1=Shingo |last2=Kiss |first2=Enikő |last3=Kuo |first3=Alan |last4=Drula |first4=Elodie |last5=Kohler |first5=Annegret |last6=Sánchez-García |first6=Marisol |last7=Morin |first7=Emmanuelle |last8=Andreopoulos |first8=Bill |last9=Barry |first9=Kerrie W. |last10=Bonito |first10=Gregory |last11=Buée |first11=Marc |last12=Carver |first12=Akiko |last13=Chen |first13=Cindy |last14=Cichocki |first14=Nicolas |last15=Clum |first15=Alicia |display-authors=3 |date=2020 |title=Large-scale genome sequencing of mycorrhizal fungi provides insights into the early evolution of symbiotic traits |journal=Nature Communications |volume=11 |issue=1 |pages=5125 |doi=10.1038/s41467-020-18795-w |pmc=7550596 |pmid=33046698 |bibcode=2020NatCo..11.5125M}}</ref> There is genetic evidence that the symbiosis between [[legume]]s and [[nitrogen-fixing bacteria]] is an extension of mycorrhizal symbiosis.<ref>{{Cite journal |last1=Provorov |first1=N. A. |last2=Shtark |first2=O. Yu |last3=Dolgikh |first3=E. A. |date=2016 |title=[Evolution of nitrogen-fixing symbioses based on the migration of bacteria from mycorrhizal fungi and soil into the plant tissues] |url=https://pubmed.ncbi.nlm.nih.gov/30024143 |journal=Zhurnal Obshchei Biologii |volume=77 |issue=5 |pages=329–345 |pmid=30024143}}</ref> The modern distribution of mycorrhizal fungi appears to reflect an increasing complexity and competition in root morphology associated with the dominance of angiosperms in the [[Cenozoic |Cenozoic Era]], characterized by complex ecological dynamics between species.<ref>{{Cite journal |last1=Brundrett |first1=Mark C. |last2=Tedersoo |first2=Leho |date=2018 |title=Evolutionary history of mycorrhizal symbioses and global host plant diversity |journal=New Phytologist  |volume=220 |issue=4 |pages=1108–1115 |doi=10.1111/nph.14976 |pmid=29355963 |doi-access=free |bibcode=2018NewPh.220.1108B }}</ref>
+===Emergence alongside terrestrial plants===
+Fossil and genetic evidence indicate that mycorrhizae emerged as early as 450-500 million years ago, potentially between fungus-like protists and algae. Arbuscular mycorrhizal relationships appeared earliest, coinciding with the [[Timeline of plant evolution#Ordovician flora|terrestrialization of plants]].<ref name="Evolution of Mycorrhiza Systems">{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |pages=467–475 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref> Genetic evidence indicates that all land plants share a single common ancestor,<ref>{{Cite journal |last1=Harris |first1=Brogan J. |last2=Clark |first2=James W. |last3=Schrempf |first3=Dominik |last4=Szöllősi |first4=Gergely J. |last5=Donoghue |first5=Philip C. J. |last6=Hetherington |first6=Alistair M. |last7=Williams |first7=Tom A. |date=2022-09-29 |title=Divergent evolutionary trajectories of bryophytes and tracheophytes from a complex common ancestor of land plants |journal=Nature Ecology & Evolution |volume=6 |issue=11 |pages=1634–1643 |doi=10.1038/s41559-022-01885-x |pmc=9630106 |pmid=36175544|bibcode=2022NatEE...6.1634H }}</ref> which appears to have quickly adopted mycorrhizal symbiosis, and research suggests that proto-mycorrhizal fungi were a key factor enabling plant terrestrialization.<ref>{{Cite journal |last1=Puginier |first1=Camille |last2=Keller |first2=Jean |last3=Delaux |first3=Pierre-Marc |date=2022-08-29 |title=Plant–microbe interactions that have impacted plant terrestrializations |url=https://academic.oup.com/plphys/article/190/1/72/6596610 |journal=Plant Physiology |volume=190 |issue=1 |pages=72–84 |doi=10.1093/plphys/kiac258 |pmid=35642902 |pmc=9434271 }}</ref> There is a strong consensus among paleomycologists that mycorrhizal fungi served as a primitive root system for early terrestrial plants.<ref name="Evolution of Mycorrhiza Systems"/> This is because, prior to plant colonization of land, soils were nutrient sparse and plants had yet to develop root systems. Without complex root systems, early terrestrial plants would have been incapable of absorbing recalcitrant ions from mineral substrates, such as phosphate, a key nutrient for plant growth.<ref name="journals.uchicago.edu">{{cite journal |last1=Maherali |first1=Hafiz |last2=Oberle |first2=Brad |last3=Stevens |first3=Peter F. |last4=Cornwell |first4=William K. |last5=McGlinn |first5=Daniel J. |title=Mutualism Persistence and Abandoment during the Evolution of the Mycorrhizal Synbiosis |journal=The American Naturalist |date=November 2016 |volume=188 |issue=5 |page=E114 |doi=10.1086/688675 |pmid=27788343 |bibcode=2016ANat..188E.113M |url=https://www.journals.uchicago.edu/doi/10.1086/688675|url-access=subscription }}</ref>
-Mycorrhizal relationships were likely crucial in terrestrial plant colonization some 450-500 million years ago, suggesting that mycorrhizal relationships are coincident with the evolution of terrestrial flora.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |pages=467–475 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>  Mycorrhizal relationships have independently evolved from saprotrophic fungi a number of times, and in effect mycorrhizae have developed multiple modes of exchange between root cells and hyphae. There are three major forms of mycorrhizal relationships which have evolved independently of one another, the oldest being arbuscular mycorrhizae, followed by ectomycorrhizal relationships, and most recently ericoid mycorrhizal relationships.
+===Fossil record and genomic analysis===
+Fossils of Glomeromycotan spores and hyphae date to 460 million years ago, but the fossils were not associated with plants. The earliest terrestrial communities were similar to modern [[Biological soil crust|biocrusts]]. Lichen-like associations between fungi and cyanobacteria were an important part of these communities. The first land plants were similar to [[moss]]es, with simple vascular systems and no leaves or roots.<ref name="nph.onlinelibrary.wiley.com">{{cite journal |last1=Strullu-Derrien |first1=Christine |last2=Selosse |first2=Marc-André |last3=Kendrick |first3=Paul |last4=Martin |first4=Francis M. |title=The Origin and Evolution of Mycorrhizal Symbioses: from Paleomycology to Phylogenomics |journal=New Phytologist |date=14 January 2018 |volume=220 |issue=4 |page=1017 |doi=10.1111/nph.15076 |pmid=29573278 |bibcode=2018NewPh.220.1012S |osti=1429513 |url=https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.15076|url-access=subscription }}</ref>
-'''Arbuscular Mycorrhizae'''
+The earliest direct fossil evidence of early mycorrhizal symbiosis is found in the 407 million year old [[Rhynie chert]], which contains an assemblage of "exceptionally preserved" fossil plants colonized by multiple para-mycorrhizal fungi.<ref name="Remy et al." /> The Rhynie chert fossils show Glomeromycotan and Mucoromycotan fungi engaged in mycorrhiza-like associations with cells of the plants [[Aglaophyton|''Aglaophyton major'']] and [[Horneophyton|''Horneophyton lignieri'']]. The fossils suggest a mutualistic association between the plants and the colonizing fungi, because distinctive nutrient exchange structures (arbuscules and hyphal coils, in Glomeromycotina and Mucoromycotina respectively) are preserved and the colonized cells appear to have been alive at the time of infection by the fungus.<ref name="nph.onlinelibrary.wiley.com"/>
-Arbuscular mycorrhizae are the oldest and most frequent form of mycorrhizal relationship.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=468 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref> Arbuscular mycorrhizae establish nutrient exchange through penetrating the root cortical cells of the host plant, making the relationship endomycorrhizal (inside the cell) as opposed to the later developed ectomycorrhizae (external nutrient exchange). Arbuscular mycorrhizae leave behind arbuscules, tree-like structures formed through hyphal penetration into the cell. Arbuscular mycorrhizae take on most angiosperms, some gymnosperms, pteridophytes, and nonvascular plants as plant hosts.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=475 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>
+These early associations are referred to as mycorrhiza-like or para-mycorrhizal because mycorrhiza are defined by the fungal colonization of plant roots, and early plants did not have any roots. The earliest fossils of arbuscular mycorrhizal fungi in plant roots originate from 315-303 million years ago, and show fungi belonging to Glomeromycotina in the root systems of a giant lycophyte, [[Lepidodendron]], and an early relative of the conifers, [[Cordaites]].<ref name="nph.onlinelibrary.wiley.com"/> Arbuscular mycorrhizal fungi were found in 240 million year old fossils of [[Antarcticycas|''Antarcticycas schopfii'']].
-Arbuscular mycorrhizas likely evolved alongside terrestrial plants approximately 450-500 million years ago when plants first began to colonize land.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=468 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>  Some scholars suggest arbuscular mycorrhizal relationships originated between fungus-like protists and algae during this time.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |pages=467–475 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref> Paramycorrhizae, mycorrhiza-like structures, have been observed in the Rhynie Chert, a 407 million-year-old piece of fossilized earth found in Scotland,<ref>{{cite journal |last1=Strullu-Derrien |first1=Christine |last2=Selosse |first2=Marc-André |last3=Kendrick |first3=Paul |last4=Martin |first4=Francis M. |title=The Origin and Evolution of Mycorrhizal Symbioses: from Paleomycology to Phylogenomics |journal=New Phytologist |date=14 January 2018 |volume=220 |issue=4 |page=1018 |doi=10.1111/nph.15076 |bibcode=2018NewPh.220.1012S |osti=1429513 |url=https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.15076|url-access=subscription }}</ref> setting a lower bound for mycorrhizal relationships. The earliest root-confined arbuscular mycorrhizae observed come from a fossil where hyphae are seen colonizing the rootlet of an arborescent clubmoss, forming arbuscules.<ref>{{cite journal |last1=Strullu-Derrien |first1=Christine |last2=Selosse |first2=Marc-André |last3=Kendrick |first3=Paul |last4=Martin |first4=Francis M. |title=The Origin and Evolution of Mycorrhizal Symbioses: from Paleomycology to Phylogenomics |journal=New Phytologist |date=14 January 2018 |volume=220 |issue=4 |page=1017 |doi=10.1111/nph.15076 |pmid=29573278 |bibcode=2018NewPh.220.1012S |osti=1429513 |url=https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.15076|url-access=subscription }}</ref>
+Ectomycorrhizae developed substantially later, during the [[Jurassic]] period, while most other modern forms of mycorrhizal symbiosis, including orchid and ericoid mycorrhizae, date to the period of [[Flowering plant#Cretaceous|angiosperm radiation]] in the [[Cretaceous]] period.<ref>{{Cite journal |last1=Miyauchi |first1=Shingo |last2=Kiss |first2=Enikő |last3=Kuo |first3=Alan |last4=Drula |first4=Elodie |last5=Kohler |first5=Annegret |last6=Sánchez-García |first6=Marisol |last7=Morin |first7=Emmanuelle |last8=Andreopoulos |first8=Bill |last9=Barry |first9=Kerrie W. |last10=Bonito |first10=Gregory |last11=Buée |first11=Marc |last12=Carver |first12=Akiko |last13=Chen |first13=Cindy |last14=Cichocki |first14=Nicolas |last15=Clum |first15=Alicia |display-authors=3 |date=2020 |title=Large-scale genome sequencing of mycorrhizal fungi provides insights into the early evolution of symbiotic traits |journal=Nature Communications |volume=11 |issue=1 |page=5125 |doi=10.1038/s41467-020-18795-w |pmc=7550596 |pmid=33046698 |bibcode=2020NatCo..11.5125M}}</ref> Ectomycorrhizae appear in the fossil record 48.7 million years ago, in the Eocene, with a fossil of ectomycorrhizal fungi colonizing ''Pinus'' roots.<ref name="The Origin and Evolution of Mycorrh">{{cite journal |last1=Strullu-Derrien |first1=Christine |last2=Selosse |first2=Marc-André |last3=Kendrick |first3=Paul |last4=Martin |first4=Francis M. |title=The Origin and Evolution of Mycorrhizal Symbioses: from Paleomycology to Phylogenomics |journal=New Phytologist |date=14 January 2018 |volume=220 |issue=4 |page=1020 |doi=10.1111/nph.15076 |pmid=29573278 |bibcode=2018NewPh.220.1012S |osti=1429513 |url=https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.15076|url-access=subscription }}</ref> However, it  is believed that the first ectomycorrhizal relationships evolved in the stem group [[Pinaceae]] around the radiation of the Pinaceae crown group in the mid Jurassic, 175 million or so years ago.<ref name="The Origin and Evolution of Mycorrh"/>
-There is a strong consensus among paleomycologists that mycorrhizal fungi served as a primitive root system for early terrestrial plants. This is because, prior to plant colonization of land, soils were nutrient sparse and plants had yet to develop root systems.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |pages=467–475 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>  Without complex root systems, early terrestrial plants would have been incapable of absorbing recalcitrant ions from mineral substrates, such as phosphate, a key nutrient for plant growth.<ref>{{cite journal |last1=Maherali |first1=Hafiz |last2=Oberle |first2=Brad |last3=Stevens |first3=Peter F. |last4=Cornwell |first4=William K. |last5=McGlinn |first5=Daniel J. |title=Mutualism Persistence and Abandoment during the Evolution of the Mycorrhizal Synbiosis |journal=The American Naturalist |date=November 2016 |volume=188 |issue=5 |page=E114 |doi=10.1086/688675 |pmid=27788343 |bibcode=2016ANat..188E.113M |url=https://www.journals.uchicago.edu/doi/10.1086/688675|url-access=subscription }}</ref> There are a number of indicators that all land plants evolved from arbuscular mycorrhizal symbiosis. One strong indicator is that arbuscular mycorrhizae have been observed in the seedling stage of otherwise ectomycorrhizal partners, suggesting that arbuscular mycorrhizae may be able to infect almost any land plant given proper circumstances.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=470 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>  Arbuscular mycorrhizal symbiosis occurs between plants and fungi in the division glomeromycota, which has been observed in almost every seed plant taxonomic division, or around 67% of species.<ref>{{cite journal |last1=Maherali |first1=Hafiz |last2=Oberle |first2=Brad |last3=Stevens |first3=Peter F. |last4=Cornwell |first4=William K. |last5=McGlinn |first5=Daniel J. |title=Mutualism Persistence and Abandoment during the Evolution of the Mycorrhizal Synbiosis |journal=The American Naturalist |date=November 2016 |volume=188 |issue=5 |page=E114 |doi=10.1086/688675 |pmid=27788343 |bibcode=2016ANat..188E.113M |url=https://www.journals.uchicago.edu/doi/10.1086/688675|url-access=subscription }}</ref> As arbuscular mycorrhizae show minimal host plant specificity, and described mycorrhizae species are likely capable of forming relationships with most host plant taxa, this also suggests that terrestrial plants and arbuscular mycorrhizae evolved with one another.
+Fossils preservation of [[Ericoid mycorrhiza]]e and orchid mycorrhizae is lacking. Calibrated molecular phylogeny is used to estimate when these mycorrhizal types originated.<ref name="The Origin and Evolution of Mycorrh"/> The origins of orchid mycorrhizae are unclear, though orchids themselves are thought to have originated in the Cretaceous period. Ericoid mycorrhizae are estimated to have the most recent evolutionary origins of mycorrhizal types, evolving around 118 million years ago from free-living saprotrophic ancestors.<ref name="ReferenceA">{{cite journal |last1=Ward |first1=Elisabeth B. |last2=Duguid |first2=Marlyse C. |last3=Kuebbing |first3=Sara E. |last4=Lendemer |first4=James C. |last5=Bradford |first5=Mark A. |title=The functional role of ericoid mycorrhizal plants and fungi on carbon and nitrogen dynamics in forests |journal=New Phytologist |date=2022 |volume=235 |issue=5 |pages=1701–1718 |doi=10.1111/nph.18307 |pmid=35704030 |bibcode=2022NewPh.235.1701W }}</ref> Ericoid mycorrhizal fungi evolved from multiple lineages of fungi, primarily ascomycetes from the [[Leotiomycetes]], as well as basidiomycetes from the family [[Serendipitaceae]].<ref>{{cite journal |last1=Perotto |first1=Silvia |last2=Daghino |first2=Stefania |last3=Martino |first3=Elena |title=Ericoid mycorrhizal fungi and their genomes: another side to the mycorrhizal symbiosis? |journal=New Phytologist |date=2018 |volume=220 |issue=4 |pages=1141–1147 |doi=10.1111/nph.15218 |pmid=29851103 |bibcode=2018NewPh.220.1141P }}</ref>
-'''Ectomycorrhizae'''
+===Origins in plants===
+In plants, the genes for forming mycorrhizal symbiosis are highly conserved and originate from a common ancestor, meaning that the ability to form mycorrhizae is ancestral to all land plants.<ref name="Mycorrhizal ecology and evolution">{{cite journal |last1=van der Heijden |first1=Marcel G. A. |last2=Martin |first2=Francis M. |last3=Selosse |first3=Marc-Andre |last4=Sanders |first4=Ian R. |title=Mycorrhizal ecology and evolution: the past, the present, and the future |journal=New Phytologist |date=2015 |volume=205 |issue=4 |pages=1406–1423 |doi=10.1111/nph.13288 |bibcode=2015NewPh.205.1406V }}</ref> Non-mycorrhizal plant lineages, such as the Brassicaceae, lost the ability to form mycorrhizae at some point in their evolution.<ref>{{cite journal |last1=Vigneron |first1=Nicolas |last2=Radhakrishnan |first2=Guru V |last3=Delaux |first3=Pierre-Marc |title=What have we learnt from studying the evolution of the arbuscular mycorrhizal symbiosis? |journal=Current Opinion in Plant Biology |date=2018 |volume=44 |pages=49–56 |doi=10.1016/j.pbi.2018.02.004 |pmid=29510317 |bibcode=2018COPB...44...49V }}</ref> The earliest  mycorrhizae were arbuscular mycorrhizae, and other forms, such as ectomycorrhizae and orchid mycorrhizae, evolved when plant hosts switched from symbiosis with [[Glomeromycotina]] to symbiosis with different fungal lineages.<ref name="At the core of the endomycorrhizal">{{cite journal |last1=Perotto |first1=Silva |last2=Balestrini |first2=Raffaella |title=At the core of the endomycorrhizal symbioses: intracellular fungal structures in orchid and arbuscular mycorrhiza |journal=New Phytologist |date=2023 |volume=242 |issue=4 |pages=1408–1416 |doi=10.1111/nph.19338 |pmid=37884478 }}</ref>
-Ectomycorrhizae are mycorrhizal relationships formed without the hyphae of the fungi penetrating the root cells of the host plant, instead forming a sheath around the root of the symbiont for nutrient exchange. The earliest confirmed ectomycorrhizal fossil dates back to the eocene approximately 48 million years ago,<ref>{{cite journal |last1=Strullu-Derrien |first1=Christine |last2=Selosse |first2=Marc-André |last3=Kendrick |first3=Paul |last4=Martin |first4=Francis M. |title=The Origin and Evolution of Mycorrhizal Symbioses: from Paleomycology to Phylogenomics |journal=New Phytologist |date=14 January 2018 |volume=220 |issue=4 |page=1020 |doi=10.1111/nph.15076 |pmid=29573278 |bibcode=2018NewPh.220.1012S |osti=1429513 |url=https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.15076|url-access=subscription }}</ref> However it’s believed that the first ectomycorrhizal relationships evolved in the stem group Pinaceae around the radiation of the Pinaceae crown group in the mid Jurassic, 175 million or so years ago. <ref>{{cite journal |last1=Strullu-Derrien |first1=Christine |last2=Selosse |first2=Marc-André |last3=Kendrick |first3=Paul |last4=Martin |first4=Francis M. |title=The Origin and Evolution of Mycorrhizal Symbioses: from Paleomycology to Phylogenomics |journal=New Phytologist |date=14 January 2018 |volume=220 |issue=4 |page=1020 |doi=10.1111/nph.15076 |pmid=29573278 |bibcode=2018NewPh.220.1012S |osti=1429513 |url=https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.15076|url-access=subscription }}</ref>
+There is genetic evidence that the symbiosis between [[legume]]s and [[nitrogen-fixing bacteria]] is derived from mycorrhizal symbiosis.<ref>{{Cite journal |last1=Provorov |first1=N. A. |last2=Shtark |first2=O. Yu |last3=Dolgikh |first3=E. A. |date=2016 |title=[Evolution of nitrogen-fixing symbioses based on the migration of bacteria from mycorrhizal fungi and soil into the plant tissues] |journal=Zhurnal Obshchei Biologii |volume=77 |issue=5 |pages=329–345 |pmid=30024143}}</ref> The modern distribution of mycorrhizal fungi appears to reflect an increasing complexity and competition in root morphology associated with the dominance of angiosperms in the [[Cenozoic |Cenozoic Era]], characterized by complex ecological dynamics between species.<ref>{{Cite journal |last1=Brundrett |first1=Mark C. |last2=Tedersoo |first2=Leho |date=2018 |title=Evolutionary history of mycorrhizal symbioses and global host plant diversity |journal=New Phytologist  |volume=220 |issue=4 |pages=1108–1115 |doi=10.1111/nph.14976 |pmid=29355963 |doi-access=free |bibcode=2018NewPh.220.1108B }}</ref>
-Ectomycorrhizal relationships have evolved a number of times, in both plants and fungi. In angiosperms, it is believed that ectomycorrhizal partnerships have evolved independently at least 18 times, and in fungi 78-82 times.<ref>{{cite journal |last1=Strullu-Derrien |first1=Christine |last2=Selosse |first2=Marc-André |last3=Kendrick |first3=Paul |last4=Martin |first4=Francis M. |title=The Origin and Evolution of Mycorrhizal Symbioses: from Paleomycology to Phylogenomics |journal=New Phytologist |date=14 January 2018 |volume=220 |issue=4 |page=1020 |doi=10.1111/nph.15076 |pmid=29573278 |bibcode=2018NewPh.220.1012S |osti=1429513 |url=https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.15076|url-access=subscription }}</ref> The main evolutionary driver for ectomycorrhizae is switching of nutritional modes from saprotrophs.<ref>{{cite journal |last1=Strullu-Derrien |first1=Christine |last2=Selosse |first2=Marc-André |last3=Kendrick |first3=Paul |last4=Martin |first4=Francis M. |title=The Origin and Evolution of Mycorrhizal Symbioses: from Paleomycology to Phylogenomics |journal=New Phytologist |date=14 January 2018 |volume=220 |issue=4 |page=1022 |doi=10.1111/nph.15076 |pmid=29573278 |bibcode=2018NewPh.220.1012S |osti=1429513 |url=https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.15076|url-access=subscription }}</ref> Phylogenomic analysis of various ectomycorrhizal fungal genomes has confirmed the
+===Origins in fungi===
-convergent evolution of ectomycorrhizal fungi from white and brown-rot fungi, as well as from soil saprotrophs – Ectomycorrhizal fungi likely evolved convergently from saprotrophic origins several times.<ref>{{cite journal |last1=Strullu-Derrien |first1=Christine |last2=Selosse |first2=Marc-André |last3=Kendrick |first3=Paul |last4=Martin |first4=Francis M. |title=The Origin and Evolution of Mycorrhizal Symbioses: from Paleomycology to Phylogenomics |journal=New Phytologist |date=14 January 2018 |volume=220 |issue=4 |page=1022 |doi=10.1111/nph.15076 |pmid=29573278 |bibcode=2018NewPh.220.1012S |osti=1429513 |url=https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.15076|url-access=subscription }}</ref><ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=470 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>  Some lineages of ectomycorrhizae have likely evolved from endophytic ancestors, fungi that live within plants without damaging them, while others such as Amanitaceae evolved from saprotrophs.<ref>{{cite journal |last1=Strullu-Derrien |first1=Christine |last2=Selosse |first2=Marc-André |last3=Kendrick |first3=Paul |last4=Martin |first4=Francis M. |title=The Origin and Evolution of Mycorrhizal Symbioses: from Paleomycology to Phylogenomics |journal=New Phytologist |date=14 January 2018 |volume=220 |issue=4 |page=1022 |doi=10.1111/nph.15076 |pmid=29573278 |bibcode=2018NewPh.220.1012S |osti=1429513 |url=https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.15076|url-access=subscription }}</ref> Some ectomycorrhizal fungi have gone through apparent evolutionary reversal back into saprotrophic ecology. This is possible because ectomycorrhizal fungi retain enzymes for breaking down lignin.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=471 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>  Most ectomycorrhizal relationships are formed between basidiomycetes or ascomycetes and woody trees or shrubs.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=469 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>
+In fungi, mycorrhizal symbiosis had multiple independent origins among different lineages of fungi. Arbuscular mycorrhizal fungi form their own monophyletic phylum, whereas other mycorrhizal fungi convergently evolved similar lifestyles.
+====Arbuscular mycorrhizae====
+The phylum [[Glomeromycota]], which forms the arbuscular mycorrhizal symbiosis, is the oldest mycorrhizal lineage. The arbuscular mycorrhizal symbiosis evolved only once in fungi; all arbuscular mycorrhizal fungi belong to Glomeromycota and share a common ancestor.<ref name="ReferenceA"/> 244 species have been identified based on differences in the appearance of their spores, but genetic studies suggest that 300-1600 species may exist in Glomeromycota.<ref name="Mycorrhizal ecology and evolution"/> All members of Glomeromycota are obligate biotrophs, entirely dependent upon their plant hosts for survival.<ref>{{cite journal |last1=Tang |first1=Nianwu |last2=San Clemente |first2=Helene |last3=Roy |first3=Sebastien |last4=Becard |first4=Guillaume |last5=Zhao |first5=Bin |last6=Roux |first6=Christophe |title=A Survey of the Gene Repertoire of Gigaspora rosea Unravels Conserved Features among Glomeromycota for Obligate Biotrophy |journal=Frontiers in Microbiology |date=2016 |volume=7 |doi=10.3389/fmicb.2016.00233 |doi-access=free }}</ref> Arbuscular mycorrhizal fungi are considered to be generalists, with minimal host plant specificity. AM symbiosis has been observed in almost every seed plant taxonomic division, or around 67% of species.<ref name="journals.uchicago.edu"/> Arbuscular mycorrhizae take on most angiosperms, some gymnosperms, pteridophytes, and nonvascular plants as plant hosts.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=475 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref> Arbuscular mycorrhizae have been observed in the seedling stage of otherwise ectomycorrhizal partners, suggesting that arbuscular mycorrhizal fungi may be able to infect almost any land plant given proper circumstances.<ref name="ReferenceB">{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=470 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>
-'''Ericoid Mycorrhizae'''
+Other forms of mycorrhizal symbiosis, such as ectomycorrhizae, orchid mycorrhizae, and ericoid mycorrhizae, emerged multiple times in different lineages of fungi through [[convergent evolution]]. Unlike arbuscular mycorrhizal fungi, some of these fungi are only facultatively symbiotic, and can live by themselves without a plant host under some conditions.
-Ericoid mycorrhizae evolved from a monophyletic origin around 140 million years ago.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=471 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>  The earliest ericoid mycorrhizae evolved from saprotrophic ascomycetes.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=471 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>  Ericoid mycorrhizae are only present in the Ericales order for plant hosts, and the Leotiales order of fungi.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=473 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>  This specialization suggests that ericoid mycorrhizal partners evolved in parallel with one another in response to environmental change, rather than through reciprocal species-to-species level selection.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=473 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>
+====Ectomycorrhizae====
+Ectomycorrhizal fungi evolved from free-living saprotrophs, mostly in [[Basidiomycota]] and [[Ascomycota]], and some became dependent on plant hosts when they lost genes necessary for decaying lignin and other plant materials.<ref name="Mycorrhizal ecology and evolution"/> There are 20,000 to 25,000 species of ectomycorrhizal fungi, but only 6,000 to 7,000 plant species that form ectomycorrhizal symbiosis.<ref>{{cite journal |last1=Voller |first1=Fay |last2=Ardanuy |first2=Agnes |last3=Taylor |first3=Andy F.S. |last4=Johnson |first4=David |title=Maintenance of host specialisation gradients in ectomycorrhizal symbionts |journal=New Phytologist |date=2023 |volume=242 |issue=4 |pages=1426–1435 |doi=10.1111/nph.19395 |pmid=37984824 }}</ref> In angiosperms, it is believed that ectomycorrhizal partnerships have developed independently at least 18 times, and in fungi, around 80 times.<ref name="The Origin and Evolution of Mycorrh"/><ref name="ReferenceA"/> The main evolutionary driver for ectomycorrhizae is switching of nutritional modes from saprotrophs.<ref name="ReferenceC">{{cite journal |last1=Strullu-Derrien |first1=Christine |last2=Selosse |first2=Marc-André |last3=Kendrick |first3=Paul |last4=Martin |first4=Francis M. |title=The Origin and Evolution of Mycorrhizal Symbioses: from Paleomycology to Phylogenomics |journal=New Phytologist |date=14 January 2018 |volume=220 |issue=4 |page=1022 |doi=10.1111/nph.15076 |pmid=29573278 |bibcode=2018NewPh.220.1012S |osti=1429513 |url=https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.15076|url-access=subscription }}</ref> Phylogenomic analysis of various ectomycorrhizal fungal genomes has confirmed the convergent evolution of ectomycorrhizal fungi from white and brown-rot fungi, as well as from soil saprotrophs.<ref name="ReferenceC"/><ref name="ReferenceB"/> Some lineages of ectomycorrhizae have likely evolved from endophytic ancestors, fungi that live within plants without damaging them.<ref name="ReferenceC"/> Some ectomycorrhizal fungi have gone through apparent evolutionary reversal back into saprotrophic ecology. This is possible because some lineages of ectomycorrhizal fungi retain enzymes for breaking down [[lignin]].<ref name="ReferenceD">{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=471 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>
-Ericoid mycorrhizal relationships are found in extremely nutrient poor soils in the northern and southern hemispheres.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=471 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>  These environments of low mineral nutrient availability have led to native plants developing sclerophylly, where plants become high in lignin and low in phosphorus and nitrogen.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=471 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>  As a result, decaying plant matter in these areas has an abnormally high carbon to nitrogen ratio, making it resistant to microbial decay. Ericoid mycorrhizae have apparently evolved to conserve minerals in nutrient deficient sclerophyllous litter by directly cycling these nutrients throughout the mycorrhiza system.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=471 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref> Ericoid mycorrhizae also retain saprotrophic abilities, allowing them to extract nitrogen and phosphorus from unmineralized organic material, and resist negative outcomes from high concentrations of toxic cations in the acidic soil environment.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=471 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref>
+====Orchid mycorrhizae====
+Orchid mycorrhizal fungi, which mostly originate from Ascomycota and Basidiomycota, are less understood. Some fungi that participate in orchid mycorrhizal symbiosis can also form ectomycorrhizal symbiosis with other plants, or live independently of a plant host. Some orchid mycorrhizal fungi can also live as plant pathogens.<ref name="At the core of the endomycorrhizal"/><ref name="Mycorrhizal ecology and evolution"/>
+====Ericoid mycorrhizae====
+Ericoid mycorrhizal associations have the most recent origins and the lowest species richness among both plant and fungal partners.<ref name="ReferenceA"/> This specialization suggests that ericoid mycorrhizal partners evolved in parallel with one another in response to environmental change, rather than through reciprocal species-to-species level selection.<ref>{{cite journal |last1=Cairney |first1=J.W.G. |title=Evolution of Mycorrhiza Systems |journal=Naturwissenschaften |date=December 2000 |volume=87 |issue=11 |page=473 |doi=10.1007/s001140050762 |pmid=11151665 |bibcode=2000NW.....87..467C }}</ref> Ericoid mycorrhizal relationships are found in extremely nutrient poor soils in the northern and southern hemispheres.<ref name="ReferenceD"/>  These environments of low mineral nutrient availability have led to native plants developing sclerophylly, where plants become high in lignin and low in phosphorus and nitrogen.<ref name="ReferenceD"/>  As a result, decaying plant matter in these areas has an abnormally high carbon to nitrogen ratio, making it resistant to microbial decay. Ericoid mycorrhizae have apparently evolved to conserve minerals in nutrient deficient sclerophyllous litter by directly cycling these nutrients throughout the mycorrhiza system.<ref name="ReferenceD"/> Ericoid mycorrhizae also retain saprotrophic abilities, allowing them to extract nitrogen and phosphorus from unmineralized organic material, and resist negative outcomes from high concentrations of toxic cations in the acidic soil environment.<ref name="ReferenceD"/>
 == Types==
-The mycorrhizal lifestyle has independently [[convergent evolution|convergently evolved]] multiple times in the history of Earth.<ref name="Ericoid mycorrhizal fungi and their">{{cite journal |last1=Perotto |first1=Silvia |last2=Daghino |first2=Stefania |last3=Martino |first3=Elena |title=Ericoid mycorrhizal fungi and their genomes: another side to the mycorrhizal symbiosis? |journal=New Phytologist |date=2018 |volume=220 |issue=4|page=1141 |doi=10.1111/nph.15218 |pmid=29851103 |bibcode=2018NewPh.220.1141P }}</ref> There are multiple ways to categorize mycorrhizal symbiosis. One major categorization is the division between ''ectomycorrhizas'' and ''endomycorrhizas''. The two types are differentiated by the fact that the hyphae of ectomycorrhizal fungi do not penetrate individual [[cell (biology)|cells]] within the root, while the [[hypha]]e of endomycorrhizal fungi penetrate the cell wall and [[wikt:invaginate|invaginate]] the [[cell membrane]].<ref>Harley, J. L.; Smith, S. E. 1983. Mycorrhizal symbiosis (1st ed.). Academic Press, London.</ref><ref name="Allen, Michael F 1991">Allen, Michael F. 1991. The ecology of mycorrhizae. Cambridge University Press, Cambridge.</ref>
+The mycorrhizal lifestyle has independently [[convergent evolution|convergently evolved]] multiple times in the history of Earth.<ref name="Ericoid mycorrhizal fungi and their">{{cite journal |last1=Perotto |first1=Silvia |last2=Daghino |first2=Stefania |last3=Martino |first3=Elena |title=Ericoid mycorrhizal fungi and their genomes: another side to the mycorrhizal symbiosis? |journal=New Phytologist |date=2018 |volume=220 |issue=4|page=1141 |doi=10.1111/nph.15218 |pmid=29851103 |bibcode=2018NewPh.220.1141P }}</ref> There are multiple ways to categorize mycorrhizal symbiosis. The largest division is between ''ectomycorrhizas'' and ''endomycorrhizas''. The two types are differentiated by the fact that the hyphae of ectomycorrhizal fungi do not penetrate individual [[cell (biology)|cells]] within the root, while the [[hypha]]e of endomycorrhizal fungi penetrate the cell wall and [[wikt:invaginate|invaginate]] the [[cell membrane]].<ref>Harley, J. L.; Smith, S. E. 1983. Mycorrhizal symbiosis (1st ed.). Academic Press, London.</ref><ref name="Allen, Michael F 1991">Allen, Michael F. 1991. The ecology of mycorrhizae. Cambridge University Press, Cambridge.</ref>
 ===Similar symbiotic relationships===
@@ Line 57: / Line 67: @@
 {{Main|Ectomycorrhiza}}
-Ectomycorrhizae are distinct in that they do not penetrate into plant cells, but instead form a structure called a [[Hartig net]] that penetrates between cells.<ref name="The potential role of Mucoromycotin">{{cite journal |last1=Howard |first1=Nathan |last2=Pressel |first2=Silvia |last3=Kaye |first3=Ryan S. |last4=Daniell |first4=Tim J. |last5=Field |first5=Katie J. |title=The potential role of Mucoromycotina 'fine root endophytes' in plant nitrogen nutrition |journal=Physiologia Plantarum |date=2022 |volume=174 |issue=3|pages=e13715 |doi=10.1111/ppl.13715 |pmid=35560043 |pmc=9328347 |bibcode=2022PPlan.174E3715H }}</ref> Ectomycorrhizas consist of a hyphal sheath, or mantle, covering the root tip and the Hartig net of hyphae surrounding the plant cells within the root [[Cortex (botany)|cortex]]. In some cases the hyphae may also penetrate the plant cells, in which case the mycorrhiza is called an endomycorrhiza. Outside the root, [[ectomycorrhizal extramatrical mycelium]] forms an extensive network within the soil and [[leaf litter]]. Other forms of mycorrhizae, including arbuscular, ericoid, arbutoid, monotropoid, and orchid mycorrhizas, are considered endomycorrhizae.<ref name="Peterson et al. 2004">{{Cite book |last1=Peterson |first1=R. L. |first2=H. B. |last2=Massicotte |name-list-style=amp |first3=L. H. |last3=Melville |date=2004 |url=http://pubs.nrc-cnrc.gc.ca/eng/books/books/9780660190877.html |title=Mycorrhizas: anatomy and cell biology |publisher=National Research Council Research Press |isbn=978-0-660-19087-7 |url-status=dead |archive-url=https://web.archive.org/web/20071225163327/http://pubs.nrc-cnrc.gc.ca/eng/books/books/9780660190877.html |archive-date=2007-12-25 }}</ref>
+Ectomycorrhizae are distinct in that they do not penetrate into plant cells, but instead form a structure called a [[Hartig net]] that penetrates between cells.<ref name="The potential role of Mucoromycotin">{{cite journal |last1=Howard |first1=Nathan |last2=Pressel |first2=Silvia |last3=Kaye |first3=Ryan S. |last4=Daniell |first4=Tim J. |last5=Field |first5=Katie J. |title=The potential role of Mucoromycotina 'fine root endophytes' in plant nitrogen nutrition |journal=Physiologia Plantarum |date=2022 |volume=174 |issue=3|article-number=e13715 |doi=10.1111/ppl.13715 |pmid=35560043 |pmc=9328347 |bibcode=2022PPlan.174E3715H }}</ref> Ectomycorrhizas consist of a hyphal sheath, or mantle, covering the root tip and the Hartig net of hyphae surrounding the plant cells within the root [[Cortex (botany)|cortex]]. In some cases the hyphae may also penetrate the plant cells, in which case the mycorrhiza is called an endomycorrhiza. Outside the root, [[ectomycorrhizal extramatrical mycelium]] forms an extensive network within the soil and [[leaf litter]]. Other forms of mycorrhizae, including arbuscular, ericoid, arbutoid, monotropoid, and orchid mycorrhizas, are considered endomycorrhizae.<ref name="Peterson et al. 2004">{{Cite book |last1=Peterson |first1=R. L. |first2=H. B. |last2=Massicotte |name-list-style=amp |first3=L. H. |last3=Melville |date=2004 |url=http://pubs.nrc-cnrc.gc.ca/eng/books/books/9780660190877.html |title=Mycorrhizas: anatomy and cell biology |publisher=National Research Council Research Press |isbn=978-0-660-19087-7 |archive-url=https://web.archive.org/web/20071225163327/http://pubs.nrc-cnrc.gc.ca/eng/books/books/9780660190877.html |archive-date=2007-12-25 }}</ref>
-Ectomycorrhizas, or EcM, are symbiotic associations between the roots of around 10% of plant families, mostly woody plants including the [[Betulaceae|birch]], [[Dipterocarpaceae|dipterocarp]], [[Myrtaceae|eucalyptus]], [[Fagaceae|oak]], [[Pinaceae|pine]], and [[Rosaceae|rose]]<ref name=Wang2006/> families, [[Orchidaceae#Ecology|orchids]],<ref>{{cite web |url=http://esciencenews.com/articles/2011/07/12/orchids.and.fungi.an.unexpected.case.symbiosis |title=Orchids and fungi: An unexpected case of symbiosis |date=July 12, 2011 |publisher=American Journal of Botany |access-date=24 July 2012 |archive-url=https://web.archive.org/web/20110715013341/http://esciencenews.com/articles/2011/07/12/orchids.and.fungi.an.unexpected.case.symbiosis |archive-date=2011-07-15 |url-status=live }}</ref> and fungi belonging to the [[Basidiomycota]], [[Ascomycota]], and [[Zygomycota]]. Ectomycorrhizae associate with relatively few plant species, only about 2% of plant species on Earth, but the species they associate with are mostly trees and woody plants that are highly dominant in their ecosystems, meaning plants in ectomycorrhizal relationships make up a large proportion of plant biomass.<ref name="The functional role of ericoid myco">{{cite journal |last1=Ward |first1=Elisabeth B. |last2=Duguid |first2=Marlyse C. |last3=Kuebbing |first3=Sara E. |last4=Lendemer |first4=James C. |last5=Bradford |first5=Mark A. |title=The functional role of ericoid mycorrhizal plants and fungi on carbon and nitrogen dynamics in forests |journal=New Phytologist |date=2022 |volume=235 |issue=5|pages=1701–1718 |doi=10.1111/nph.18307 |pmid=35704030 |bibcode=2022NewPh.235.1701W }}</ref> 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.<ref name=bak04/> An individual tree may have 15 or more different fungal EcM partners at one time.<ref name=saari04/> While the diversity of plants involved in EcM is low, the diversity of fungi involved in EcM is high. 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 probably be between 20,000 and 25,000.<ref>{{cite journal |last1=Rinaldi |first1=A. C. |last2=Comandini |first2=O. |last3=Kuyper |first3=T. W. |date=2008 |title=Ectomycorrhizal fungal diversity: separating the wheat from the chaff |journal=Fungal Diversity |volume=33 |pages=1–45 |url=http://www.fungaldiversity.org/fdp/sfdp/33-1.pdf |access-date=2011-05-23 |archive-url=https://web.archive.org/web/20110724163606/http://www.fungaldiversity.org/fdp/sfdp/33-1.pdf |archive-date=2011-07-24 |url-status=live }}</ref> Ectomycorrhizal fungi evolved independently from saprotrophic ancestors many times in the group's history.<ref>{{cite journal |last1=Martin |first1=Francis M. |last2=van der Heijden |first2=Marcel G. A. |title=The mycorrhizal symbiosis: research frontiers in genomics, ecology, and agricultural application |journal=New Phytologist |date=2024 |volume=242 |issue=4|pages=1486–1506 |doi=10.1111/nph.19541 |pmid=38297461 |bibcode=2024NewPh.242.1486M }}</ref>
+Ectomycorrhizas, or EcM, are symbiotic associations between the roots of around 10% of plant families, mostly woody plants including the [[Betulaceae|birch]], [[Dipterocarpaceae|dipterocarp]], [[Myrtaceae|eucalyptus]], [[Fagaceae|oak]], [[Pinaceae|pine]], and [[Rosaceae|rose]]<ref name=Wang2006/> families, [[Orchidaceae#Ecology|orchids]],<ref>{{cite web |url=http://esciencenews.com/articles/2011/07/12/orchids.and.fungi.an.unexpected.case.symbiosis |title=Orchids and fungi: An unexpected case of symbiosis |date=July 12, 2011 |publisher=American Journal of Botany |access-date=24 July 2012 |archive-url=https://web.archive.org/web/20110715013341/http://esciencenews.com/articles/2011/07/12/orchids.and.fungi.an.unexpected.case.symbiosis |archive-date=2011-07-15 |url-status=live }}</ref> and fungi belonging to the [[Basidiomycota]], [[Ascomycota]], and [[Zygomycota]]. Ectomycorrhizae associate with relatively few plant species, only about 2% of plant species on Earth, but the species they associate with are mostly trees and woody plants that are highly dominant in their ecosystems, meaning plants in ectomycorrhizal relationships make up a large proportion of plant biomass.<ref name="The functional role of ericoid myco">{{cite journal |last1=Ward |first1=Elisabeth B. |last2=Duguid |first2=Marlyse C. |last3=Kuebbing |first3=Sara E. |last4=Lendemer |first4=James C. |last5=Bradford |first5=Mark A. |title=The functional role of ericoid mycorrhizal plants and fungi on carbon and nitrogen dynamics in forests |journal=New Phytologist |date=2022 |volume=235 |issue=5|pages=1701–1718 |doi=10.1111/nph.18307 |pmid=35704030 |bibcode=2022NewPh.235.1701W }}</ref> 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.<ref name=bak04/> An individual tree may have 15 or more different fungal EcM partners at one time.<ref name=saari04/> While the diversity of plants involved in EcM is low, the diversity of fungi involved in EcM is high. 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 probably be between 20,000 and 25,000.<ref>{{cite journal |last1=Rinaldi |first1=A. C. |last2=Comandini |first2=O. |last3=Kuyper |first3=T. W. |date=2008 |title=Ectomycorrhizal fungal diversity: separating the wheat from the chaff |journal=Fungal Diversity |volume=33 |pages=1–45 |url=http://www.fungaldiversity.org/fdp/sfdp/33-1.pdf |access-date=2011-05-23 |archive-url=https://web.archive.org/web/20110724163606/http://www.fungaldiversity.org/fdp/sfdp/33-1.pdf |archive-date=2011-07-24 |url-status=live }}</ref> Ectomycorrhizal fungi evolved independently from saprotrophic ancestors many times in the group's history.<ref name="The mycorrhizal symbiosis: research">{{cite journal |last1=Martin |first1=Francis M. |last2=van der Heijden |first2=Marcel G. A. |title=The mycorrhizal symbiosis: research frontiers in genomics, ecology, and agricultural application |journal=New Phytologist |date=2024 |volume=242 |issue=4|pages=1486–1506 |doi=10.1111/nph.19541 |pmid=38297461 |bibcode=2024NewPh.242.1486M }}</ref>
 Nutrients can be shown to move between different plants through the fungal network. Carbon has been shown to move from [[Betula papyrifera|paper birch]] seedlings into adjacent [[Coast Douglas-fir|Douglas-fir]] seedlings, although not conclusively through a common mycorrhizal network,<ref>{{Cite journal |last1=Karst |first1=Justine |last2=Jones |first2=Melanie D. |last3=Hoeksema |first3=Jason D. |date=2023-02-13 |title=Positive citation bias and overinterpreted results lead to misinformation on common mycorrhizal networks in forests |url=https://www.nature.com/articles/s41559-023-01986-1 |journal=Nature Ecology & Evolution |language=en |volume=7 |issue=4 |pages=501–511 |doi=10.1038/s41559-023-01986-1 |pmid=36782032 |bibcode=2023NatEE...7..501K |s2cid=256845005 |issn=2397-334X|url-access=subscription }}</ref> thereby promoting [[Ecological succession|succession]] in [[ecosystem]]s.<ref>{{cite journal |last1=Simard |first1=Suzanne W. |last2=Perry |first2=David A. |last3=Jones |first3=Melanie D. |last4=Myrold |first4=David D. |last5=Durall |first5=Daniel M. |last6=Molina |first6=Randy |name-list-style=amp |title=Net transfer of carbon between ectomycorrhizal tree species in the field |journal=Nature |volume=388 |issue=6642 |date=1997 |pages=579–582 |doi=10.1038/41557 |bibcode=1997Natur.388..579S |s2cid=4423207 |doi-access=free }}</ref> The ectomycorrhizal fungus ''[[Laccaria bicolor]]'' has been found to lure and kill [[springtail]]s 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.<ref>[https://archive.today/20120710035451/http://findarticles.com/p/articles/mi_m1200/is_14_159/ai_104730213/ Fungi kill insects and feed host plants] BNET.com</ref><ref>{{cite journal |last1=Klironomos |first1=J. N. |last2=Hart |first2=M. M. |date=2001 |title=Animal nitrogen swap for plant carbon |journal=Nature |volume=410 |issue=6829 |pages=651–652 |doi=10.1038/35070643 |pmid=11287942 |bibcode=2001Natur.410..651K |s2cid=4418192 }}</ref> When compared with non-mycorrhizal fine roots, ectomycorrhizae may contain very high concentrations of trace elements, including toxic metals (cadmium, silver) or chlorine.<ref>{{cite journal |last1=Cejpková |first1=J. |last2=Gryndler |first2=M. |last3=Hršelová |first3=H. |last4=Kotrba |first4=P. |last5=Řanda |first5=Z. |last6=Greňová |first6=I. |last7=Borovička |first7=J. |title=Bioaccumulation of heavy metals, metalloids, and chlorine in ectomycorrhizae from smelter-polluted area |journal=Environmental Pollution |volume=218 |pages=176–185 |doi=10.1016/j.envpol.2016.08.009 |pmid=27569718 |year=2016|bibcode=2016EPoll.218..176C }}</ref>
-The first genomic sequence for a representative of symbiotic fungi, the ectomycorrhizal basidiomycete ''L. bicolor'', was published in 2008.<ref>{{cite journal |last1=Martin |first1=F. |date=2008 |title=The genome of ''Laccaria bicolor'' provides insights into mycorrhizal symbiosis |doi=10.1038/nature06556 |journal=Nature |volume=452 |issue=7183 |pages=88–92 |pmid=18322534 |first2=A. |last3=Ahrén |first3=D. |last4=Brun |first4=A. |last5=Danchin |first5=E. G. J. |last6=Duchaussoy |first6=F. |last7=Gibon |first7=J. |last8=Kohler |first8=A. |last9=Lindquist |first9=E. |display-authors=2 |last2=Aerts |bibcode=2008Natur.452...88M |url=https://nootropicsfrontline.com/wp-content/uploads/2021/07/wiki_martin2008.pdf |doi-access=free }}</ref> 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. ''L. 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, ''L. 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. Since then, the genomes of many other ectomycorrhizal fungal species have been sequenced further expanding the study of gene families and evolution in these organisms.<ref>{{Cite journal |last1=Miyauchi |first1=Shingo |last2=Kiss |first2=Enikő |last3=Kuo |first3=Alan |last4=Drula |first4=Elodie |last5=Kohler |first5=Annegret |last6=Sánchez-García |first6=Marisol |last7=Morin |first7=Emmanuelle |last8=Andreopoulos |first8=Bill |last9=Barry |first9=Kerrie W. |last10=Bonito |first10=Gregory |last11=Buée |first11=Marc |last12=Carver |first12=Akiko |last13=Chen |first13=Cindy |last14=Cichocki |first14=Nicolas |last15=Clum |first15=Alicia |date=2020-10-12 |title=Large-scale genome sequencing of mycorrhizal fungi provides insights into the early evolution of symbiotic traits |journal=Nature Communications |language=en |volume=11 |issue=1 |pages=5125 |doi=10.1038/s41467-020-18795-w |issn=2041-1723 |pmc=7550596 |pmid=33046698|bibcode=2020NatCo..11.5125M }}</ref>
+The first genomic sequence for a representative of symbiotic fungi, the ectomycorrhizal basidiomycete ''L. bicolor'', was published in 2008.<ref>{{cite journal |last1=Martin |first1=F. |date=2008 |title=The genome of ''Laccaria bicolor'' provides insights into mycorrhizal symbiosis |doi=10.1038/nature06556 |journal=Nature |volume=452 |issue=7183 |pages=88–92 |pmid=18322534 |first2=A. |last3=Ahrén |first3=D. |last4=Brun |first4=A. |last5=Danchin |first5=E. G. J. |last6=Duchaussoy |first6=F. |last7=Gibon |first7=J. |last8=Kohler |first8=A. |last9=Lindquist |first9=E. |display-authors=2 |last2=Aerts |bibcode=2008Natur.452...88M |url=https://nootropicsfrontline.com/wp-content/uploads/2021/07/wiki_martin2008.pdf |doi-access=free }}</ref> 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. ''L. 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, ''L. 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. Since then, the genomes of many other ectomycorrhizal fungal species have been sequenced further expanding the study of gene families and evolution in these organisms.<ref>{{Cite journal |last1=Miyauchi |first1=Shingo |last2=Kiss |first2=Enikő |last3=Kuo |first3=Alan |last4=Drula |first4=Elodie |last5=Kohler |first5=Annegret |last6=Sánchez-García |first6=Marisol |last7=Morin |first7=Emmanuelle |last8=Andreopoulos |first8=Bill |last9=Barry |first9=Kerrie W. |last10=Bonito |first10=Gregory |last11=Buée |first11=Marc |last12=Carver |first12=Akiko |last13=Chen |first13=Cindy |last14=Cichocki |first14=Nicolas |last15=Clum |first15=Alicia |date=2020-10-12 |title=Large-scale genome sequencing of mycorrhizal fungi provides insights into the early evolution of symbiotic traits |journal=Nature Communications |language=en |volume=11 |issue=1 |page=5125 |doi=10.1038/s41467-020-18795-w |issn=2041-1723 |pmc=7550596 |pmid=33046698|bibcode=2020NatCo..11.5125M }}</ref>
 ====Arbutoid mycorrhiza====
@@ Line 80: / Line 90: @@
 ===Mucoromycotina fine root endophytes===
-Mycorrhizal fungi belonging to [[Mucoromycotina]], known as “fine root endophytes" (MFREs), were mistakenly identified as arbuscular mycorrhizal fungi until recently. While similar to AMF, MFREs are from subphylum Mucoromycotina instead of Glomeromycotina. Their morphology when colonizing a plant root is very similar to AMF, but they form fine textured hyphae.<ref name="The potential role of Mucoromycotin"/> Effects of MFREs may have been mistakenly attributed to AMFs due to confusion between the two, complicated by the fact that AMFs and MFREs often colonize the same hosts simultaneously. Unlike AMFs, they appear capable of surviving without a host. This group of mycorrhizal fungi is little understood, but appears to prefer wet, acidic soils and forms symbiotic relationships with liverworts, hornworts, lycophytes, and angiosperms.<ref>{{cite journal |last1=Prout |first1=James N. |last2=Williams |first2=Alex |last3=Wanke |first3=Alan |last4=Schornack |first4=Sebastian |last5=Ton |first5=Jurriaan |last6=Field |first6=Katie J. |title=Mucoromycotina 'fine root endophytes': a new molecular model for plant–fungal mutualisms? |journal=Trends in Plant Science |date=2023 |volume=29 |issue=6|pmid=38102045 }}</ref>
+Mycorrhizal fungi belonging to [[Mucoromycotina]], known as "fine root endophytes" (MFREs), were mistakenly identified as arbuscular mycorrhizal fungi until recently. While similar to AMF, MFREs are from subphylum Mucoromycotina instead of Glomeromycotina. Their morphology when colonizing a plant root is very similar to AMF, but they form fine textured hyphae.<ref name="The potential role of Mucoromycotin"/> Effects of MFREs may have been mistakenly attributed to AMFs due to confusion between the two, complicated by the fact that AMFs and MFREs often colonize the same hosts simultaneously. Unlike AMFs, they appear capable of surviving without a host. This group of mycorrhizal fungi is little understood, but appears to prefer wet, acidic soils and forms symbiotic relationships with liverworts, hornworts, lycophytes, and angiosperms.<ref name="Mucoromycotina 'fine root endophyte">{{cite journal |last1=Prout |first1=James N. |last2=Williams |first2=Alex |last3=Wanke |first3=Alan |last4=Schornack |first4=Sebastian |last5=Ton |first5=Jurriaan |last6=Field |first6=Katie J. |title=Mucoromycotina 'fine root endophytes': a new molecular model for plant–fungal mutualisms? |journal=Trends in Plant Science |date=2023 |volume=29 |issue=6|pmid=38102045 }}</ref>
 ===Ericoid mycorrhiza===
@@ Line 98: / Line 108: @@
 {{Main |Orchid mycorrhiza}}
-All [[Orchidaceae|orchids]] are [[myco-heterotrophy|myco-heterotrophic]] at some stage during their lifecycle, meaning that they can survive only if they form [[orchid mycorrhiza]]e. Orchid seeds are so small that they contain no nutrition to sustain the germinating seedling, and instead must gain the energy to grow from their fungal symbiont.<ref name="The potential role of Mucoromycotin"/> The OM relationship is asymmetric; the plant seems to benefit more than the fungus, and some orchids are entirely mycoheterotrophic, lacking chlorophyll for photosynthesis. It is actually unknown whether fully autotrophic orchids that do not receive some of their carbon from fungi exist or not.<ref>{{cite journal |last1=Li |first1=Taiqiang |last2=Yang |first2=Wenke |last3=Wu |first3=Shimao |last4=Selosse |first4=Marc-Andre |last5=Gao |first5=Jiangyun |title=Progress and Prospects of Mycorrhizal Fungal Diversity in Orchids |journal=Frontiers in Plant Science |date=2021 |volume=12|doi=10.3389/fpls.2021.646325 |doi-access=free |pmid=34025694 |pmc=8138444 |bibcode=2021FrPS...1246325L }}</ref> Like fungi that form ErMs, OM fungi can sometimes live as endophytes or as independent saprotrophs. In the OM symbiosis, hyphae penetrate into the root cells and form pelotons (coils) for nutrient exchange.
+All [[Orchidaceae|orchids]] are [[myco-heterotrophy|myco-heterotrophic]] at some stage during their lifecycle, meaning that they can survive only if they form [[orchid mycorrhiza]]e. Orchid seeds are so small that they contain no nutrition to sustain the germinating seedling, and instead must gain the energy to grow from their fungal symbiont.<ref name="The potential role of Mucoromycotin"/> The OM relationship is asymmetric; the plant seems to benefit more than the fungus, and some orchids are entirely mycoheterotrophic, lacking chlorophyll for photosynthesis. It is actually unknown whether fully autotrophic orchids that do not receive some of their carbon from fungi exist or not.<ref>{{cite journal |last1=Li |first1=Taiqiang |last2=Yang |first2=Wenke |last3=Wu |first3=Shimao |last4=Selosse |first4=Marc-Andre |last5=Gao |first5=Jiangyun |title=Progress and Prospects of Mycorrhizal Fungal Diversity in Orchids |journal=Frontiers in Plant Science |date=2021 |volume=12|article-number=646325 |doi=10.3389/fpls.2021.646325 |doi-access=free |pmid=34025694 |pmc=8138444 |bibcode=2021FrPS...1246325L }}</ref> Like fungi that form ErMs, OM fungi can sometimes live as endophytes or as independent saprotrophs. In the OM symbiosis, hyphae penetrate into the root cells and form pelotons (coils) for nutrient exchange.
 ===Monotropoid mycorrhiza===
+[[File:Indian pipe PDB.JPG|thumb|alt='Monotropa' plant unable to photosynthesis, collects food from monotropoid mycorrhiza|[[Monotropa uniflora|''Monotropa'']] plant unable to photosynthesis, collects food from monotropoid mycorrhiza]]
 {{Main |Myco-heterotrophy}}
@@ Line 117: / Line 127: @@
 ===Formation===
-To successfully engage in mutualistic symbiotic relationships with [[Endophyte|other organisms]], such as mycorrhizal fungi and any of the thousands of microbes that colonize plants, plants must discriminate between mutualists and pathogens, allowing the mutualists to colonize while activating an [[Immune system|immune]] response towards the pathogens. Plant genomes code for potentially hundreds of [[Receptor (biochemistry)|receptors]] for detecting chemical signals from other organisms. Plants dynamically adjust their symbiotic and immune responses, changing their interactions with their symbionts in response to feedbacks detected by the plant.<ref>{{cite journal |last1=Thoms |first1=David |last2=Liang |first2=Yan |last3=Haney |first3=Cara H. |title=Maintaining Symbiotic Homeostasis: How Do Plants Engage With Beneficial Microorganisms While at the Same Time Restricting Pathogens? |journal=International Society for Molecular Plant-Microbe Interactions |date=2021 |volume=34 |issue=5}}</ref> In plants, the mycorrhizal symbiosis is regulated by the [[Common symbiosis signaling pathway|common symbiosis signaling pathway (CSSP)]], a set of genes involved in initiating and maintaining colonization by endosymbiotic fungi and other endosymbionts such as [[Rhizobia]] in [[legume]]s. The CSSP has origins predating the colonization of land by plants, demonstrating that the co-evolution of plants and arbuscular mycorrhizal fungi is over 500 million years old.<ref>{{cite journal |last1=Martin |first1=Francis M. |last2=van der Heijden |first2=Marcel G. A. |title=The mycorrhizal symbiosis: research frontiers in genomics, ecology, and agricultural application |journal=New Phytologist |date=2024 |volume=242 |issue=4|pages=1486–1506 |doi=10.1111/nph.19541 |pmid=38297461 |bibcode=2024NewPh.242.1486M }}</ref> In arbuscular mycorrhizal fungi, the presence of [[strigolactone]]s, a plant hormone, secreted from roots induces fungal spores in the soil to germinate, stimulates their metabolism, growth and branching, and prompts the fungi to release chemical signals the plant can detect.<ref>{{cite journal |last1=Ho-Plagaro |first1=Tania |last2=Garcia-Garrido |first2=Jose Manuel |title=Molecular Regulation of Arbuscular Mycorrhizal Symbiosis |journal=International Journal of Molecular Sciences |date=2022 |volume=23 |issue=11|page=5960 |doi=10.3390/ijms23115960 |doi-access=free |pmid=35682640 |pmc=9180548 }}</ref> Once the plant and fungus recognize one another as suitable symbionts, the plant activates the common symbiotic signaling pathway, which causes changes in the root tissues that enable the fungus to colonize.<ref>{{cite journal |last1=Nasir |first1=Fahad |last2=Bahadur |first2=Ali |last3=Lin |first3=Xiaolong |last4=Gao |first4=Yingzhi |last5=Tian |first5=Chunjie |title=Novel insights into host receptors and receptor-mediated signaling that regulate arbuscular mycorrhizal symbiosis |journal=Journal of Experimental Botany |date=2021 |volume=72 |issue=5|pages=1546–1557 |doi=10.1093/jxb/eraa538 |pmid=33252650 }}</ref>
+To successfully engage in mutualistic symbiotic relationships with [[Endophyte|other organisms]], such as mycorrhizal fungi and any of the thousands of microbes that colonize plants, plants must discriminate between mutualists and pathogens, allowing the mutualists to colonize while activating an [[Immune system|immune]] response towards the pathogens. Plant genomes code for potentially hundreds of [[Receptor (biochemistry)|receptors]] for detecting chemical signals from other organisms. Plants dynamically adjust their symbiotic and immune responses, changing their interactions with their symbionts in response to feedbacks detected by the plant.<ref>{{cite journal |last1=Thoms |first1=David |last2=Liang |first2=Yan |last3=Haney |first3=Cara H. |title=Maintaining Symbiotic Homeostasis: How Do Plants Engage With Beneficial Microorganisms While at the Same Time Restricting Pathogens? |journal=International Society for Molecular Plant-Microbe Interactions |date=2021 |volume=34 |issue=5}}</ref> In plants, the mycorrhizal symbiosis is regulated by the [[Common symbiosis signaling pathway|common symbiosis signaling pathway (CSSP)]], a set of genes involved in initiating and maintaining colonization by endosymbiotic fungi and other endosymbionts such as [[Rhizobia]] in [[legume]]s. The CSSP has origins predating the colonization of land by plants, demonstrating that the co-evolution of plants and arbuscular mycorrhizal fungi is over 500 million years old.<ref name="The mycorrhizal symbiosis: research"/> In arbuscular mycorrhizal fungi, the presence of [[strigolactone]]s, a plant hormone, secreted from roots induces fungal spores in the soil to germinate, stimulates their metabolism, growth and branching, and prompts the fungi to release chemical signals the plant can detect.<ref name="Molecular Regulation of Arbuscular">{{cite journal |last1=Ho-Plagaro |first1=Tania |last2=Garcia-Garrido |first2=Jose Manuel |title=Molecular Regulation of Arbuscular Mycorrhizal Symbiosis |journal=International Journal of Molecular Sciences |date=2022 |volume=23 |issue=11|page=5960 |doi=10.3390/ijms23115960 |doi-access=free |pmid=35682640 |pmc=9180548 }}</ref> Once the plant and fungus recognize one another as suitable symbionts, the plant activates the common symbiotic signaling pathway, which causes changes in the root tissues that enable the fungus to colonize.<ref>{{cite journal |last1=Nasir |first1=Fahad |last2=Bahadur |first2=Ali |last3=Lin |first3=Xiaolong |last4=Gao |first4=Yingzhi |last5=Tian |first5=Chunjie |title=Novel insights into host receptors and receptor-mediated signaling that regulate arbuscular mycorrhizal symbiosis |journal=Journal of Experimental Botany |date=2021 |volume=72 |issue=5|pages=1546–1557 |doi=10.1093/jxb/eraa538 |pmid=33252650 }}</ref>
-Experiments with arbuscular mycorrhizal fungi have identified numerous chemical compounds to be involved in the "chemical dialog" that occurs between the prospective symbionts before symbiosis is begun. In plants, almost all plant hormones play a role in initiating or regulating AMF symbiosis, and other chemical compounds are also suspected to have a signaling function. While the signals emitted by the fungi are less understood, it has been shown that chitinaceous molecules known as Myc factors are essential for the formation of arbuscular mycorrhizae. Signals from plants are detected by LysM-containing receptor-like kinases, or LysM-RLKs. AMF genomes also code for potentially hundreds of effector proteins, of which only a few have a proven effect on mycorrhizal symbiosis, but many others likely have a function in communication with plant hosts as well.<ref>{{cite journal |last1=Ho-Plagaro |first1=Tania |last2=Garcia-Garrido |first2=Jose Manuel |title=Molecular Regulation of Arbuscular Mycorrhizal Symbiosis |journal=International Journal of Molecular Sciences |date=2022 |volume=23 |issue=11|page=5960 |doi=10.3390/ijms23115960 |doi-access=free |pmid=35682640 |pmc=9180548 }}</ref>
+Experiments with arbuscular mycorrhizal fungi have identified numerous chemical compounds to be involved in the "chemical dialog" that occurs between the prospective symbionts before symbiosis is begun. In plants, almost all plant hormones play a role in initiating or regulating AMF symbiosis, and other chemical compounds are also suspected to have a signaling function. While the signals emitted by the fungi are less understood, it has been shown that chitinaceous molecules known as Myc factors are essential for the formation of arbuscular mycorrhizae. Signals from plants are detected by LysM-containing receptor-like kinases, or LysM-RLKs. AMF genomes also code for potentially hundreds of effector proteins, of which only a few have a proven effect on mycorrhizal symbiosis, but many others likely have a function in communication with plant hosts as well.<ref name="Molecular Regulation of Arbuscular"/>
-Many factors are involved in the initiation of mycorrhizal symbiosis, but particularly influential is the plant's need for [[phosphorus]]. Experiments involving [[Oryza sativa|rice]] plants with a mutation disabling their ability to detect P starvation show that arbuscular mycorrhizal fungi detection, recruitment and colonization is prompted when the plant detects that it is starved of phosphorus.<ref>{{cite journal |last1=Prout |first1=James N. |last2=Williams |first2=Alex |last3=Wanke |first3=Alan |last4=Schornack |first4=Sebastian |last5=Ton |first5=Jurriaan |last6=Field |first6=Katie J. |title=Mucoromycotina 'fine root endophytes': a new molecular model for plant–fungal mutualisms? |journal=Trends in Plant Science |date=2023 |volume=29 |issue=6|pmid=38102045 }}</ref> Nitrogen starvation also plays a role in initiating AMF symbiosis.<ref>{{cite journal |last1=Ho-Plagaro |first1=Tania |last2=Garcia-Garrido |first2=Jose Manuel |title=Molecular Regulation of Arbuscular Mycorrhizal Symbiosis |journal=International Journal of Molecular Sciences |date=2022 |volume=23 |issue=11|page=5960 |doi=10.3390/ijms23115960 |doi-access=free |pmid=35682640 |pmc=9180548 }}</ref>
+Many factors are involved in the initiation of mycorrhizal symbiosis, but particularly influential is the plant's need for [[phosphorus]]. Experiments involving [[Oryza sativa|rice]] plants with a mutation disabling their ability to detect P starvation show that arbuscular mycorrhizal fungi detection, recruitment and colonization is prompted when the plant detects that it is starved of phosphorus.<ref name="Mucoromycotina 'fine root endophyte"/> Nitrogen starvation also plays a role in initiating AMF symbiosis.<ref name="Molecular Regulation of Arbuscular"/>
 ===Mechanisms===
-The mechanisms by which mycorrhizae increase absorption include some that are physical and some that are chemical. Physically, most mycorrhizal mycelia are much smaller in diameter than the smallest root or root hair, and thus can explore soil material that roots and root hairs cannot reach, and provide a larger surface area for absorption. Chemically, the cell membrane chemistry of fungi differs from that of plants. For example, they may secrete [[organic acid]]s that dissolve or [[Chelation|chelate]] many ions, or release them from minerals by [[ion exchange]].<ref>{{cite book |chapter=Overview of Mycorrhizal Symbioses |title=Principles and Applications of Soil Microbiology |isbn=978-0-13-094117-6 |chapter-url=http://cropsoil.psu.edu/sylvia/mycorrhiza.htm |archive-url=https://web.archive.org/web/20100623051447/http://cropsoil.psu.edu/sylvia/mycorrhiza.htm|archive-date= June 23, 2010|last1=Sylvia |first1=David M. |last2=Fuhrmann |first2=Jeffry J. |last3=Hartel |first3=Peter G. |last4=Zuberer |first4=David A. |year=2005 |publisher=Pearson Prentice Hall }}</ref> Mycorrhizae are especially beneficial for the plant partner in nutrient-poor soils.<ref>{{cite web |url=http://www.biologie.uni-hamburg.de/b-online/e33/33b.htm |title=Botany online: Interactions - Plants - Fungi - Parasitic and Symbiotic Relations - Mycorrhiza |publisher=Biologie.uni-hamburg.de |access-date=2010-09-30 |url-status=dead |archive-url=https://web.archive.org/web/20110606050759/http://www.biologie.uni-hamburg.de/b-online/e33/33b.htm |archive-date=2011-06-06 }}</ref>
+The mechanisms by which mycorrhizae increase absorption include some that are physical and some that are chemical. Physically, most mycorrhizal mycelia are much smaller in diameter than the smallest root or root hair, and thus can explore soil material that roots and root hairs cannot reach, and provide a larger surface area for absorption. Chemically, the cell membrane chemistry of fungi differs from that of plants. For example, they may secrete [[organic acid]]s that dissolve or [[Chelation|chelate]] many ions, or release them from minerals by [[ion exchange]].<ref>{{cite book |chapter=Overview of Mycorrhizal Symbioses |title=Principles and Applications of Soil Microbiology |isbn=978-0-13-094117-6 |chapter-url=http://cropsoil.psu.edu/sylvia/mycorrhiza.htm |archive-url=https://web.archive.org/web/20100623051447/http://cropsoil.psu.edu/sylvia/mycorrhiza.htm|archive-date= June 23, 2010|last1=Sylvia |first1=David M. |last2=Fuhrmann |first2=Jeffry J. |last3=Hartel |first3=Peter G. |last4=Zuberer |first4=David A. |year=2005 |publisher=Pearson Prentice Hall }}</ref> Mycorrhizae are especially beneficial for the plant partner in nutrient-poor soils.<ref>{{cite web |url=http://www.biologie.uni-hamburg.de/b-online/e33/33b.htm |title=Botany online: Interactions - Plants - Fungi - Parasitic and Symbiotic Relations - Mycorrhiza |publisher=Biologie.uni-hamburg.de |access-date=2010-09-30 |archive-url=https://web.archive.org/web/20110606050759/http://www.biologie.uni-hamburg.de/b-online/e33/33b.htm |archive-date=2011-06-06 }}</ref>
 ===Sugar-water/mineral exchange===
@@ Line 138: / Line 148: @@
 ''[[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 fixation|nitrogen fixing]] [[bacteria]] which contribute a significant amount of [[nitrogen]] and allow the pines to colonize nutrient-poor sites.<ref name=paul07/>
 ===Disease, drought and salinity resistance and its correlation to mycorrhizae===
 Mycorrhizal plants are often more resistant to diseases, such as those caused by microbial soil-borne [[pathogen]]s. These associations have been found to assist in plant defense both above and belowground. Mycorrhizas have been found to excrete enzymes that are toxic to soil borne organisms such as nematodes.<ref>{{cite journal |last1=Azcón-Aguilar |first1=C. |last2=Barea |first2=J. M. |title=Arbuscular mycorrhizas and biological control of soil-borne plant pathogens – an overview of the mechanisms involved |journal=Mycorrhiza |date=29 October 1996 |volume=6 |issue=6 |pages=457–464 |doi=10.1007/s005720050147 |s2cid=25190159}}</ref> More recent studies have shown that mycorrhizal associations result in a priming effect of plants that essentially acts as a primary immune response. When this association is formed a defense response is activated similarly to the response that occurs when the plant is under attack. As a result of this inoculation, defense responses are stronger in plants with mycorrhizal associations.<ref>{{cite journal |last1=Jung |first1=Sabine C. |last2=Martinez-Medina |first2=Ainhoa |last3=Lopez-Raez |first3=Juan A. |last4=Pozo |first4=Maria J. |title=Mycorrhiza-Induced Resistance and Priming of Plant Defenses |journal=J Chem Ecol |date=24 May 2012 |volume=38 |issue=6 |pages=651–664 |doi=10.1007/s10886-012-0134-6 |pmid=22623151 |bibcode=2012JCEco..38..651J |s2cid=12918193|hdl=10261/344431 |hdl-access=free }}</ref>
-[[Ecosystem services]] provided by mycorrhizal fungi may depend on the soil microbiome.<ref name="Svenningsen 1296–1307">{{cite journal |last1=Svenningsen |first1=Nanna B |last2=Watts-Williams |first2=Stephanie J |last3=Joner |first3=Erik J |last4=Battini |first4=Fabio |last5=Efthymiou |first5=Aikaterini |last6=Cruz-Paredes |first6=Carla |last7=Nybroe |first7=Ole |last8=Jakobsen |first8=Iver |title=Suppression of the activity of arbuscular mycorrhizal fungi by the soil microbiota |journal=The ISME Journal |date=May 2018 |volume=12 |issue=5 |pages=1296–1307 |doi=10.1038/s41396-018-0059-3 |pmid=29382946 |pmc=5931975 |doi-access=free |bibcode=2018ISMEJ..12.1296S }}</ref> Furthermore, mycorrhizal fungi was significantly correlated with soil physical variable, but only with water level and not with aggregate stability<ref>{{cite book |doi=10.1007/1-4020-4447-X_10 |chapter=Disease Resistance in Plants Through Mycorrhizal Fungi Induced Allelochemicals |title=Allelochemicals: Biological Control of Plant Pathogens and Diseases |series=Disease Management of Fruits and Vegetables |year=2006 |last1=Zeng |first1=Ren-Sen |volume=2 |pages=181–192 |isbn=1-4020-4445-3 }}</ref><ref>{{cite web |url=https://www.usask.ca/biology/kaminskyj/arctic.html |title=Dr. Susan Kaminskyj: Endorhizal Fungi |publisher=Usask.ca |access-date=2010-09-30 |archive-url=https://web.archive.org/web/20101104000757/http://www.usask.ca/biology/kaminskyj/arctic.html |archive-date=2010-11-04 |url-status=dead }}</ref> and can lead also to more resistant to the effects of drought.<ref>{{cite web |url=http://aggie-horticulture.tamu.edu/Faculty/davies/research/mycorrhizae.html |title=Dr. Davies Research Page |publisher=Aggie-horticulture.tamu.edu |access-date=2010-09-30 |url-status=dead |archive-url=https://web.archive.org/web/20101019002159/http://aggie-horticulture.tamu.edu/faculty/davies/research/mycorrhizae.html |archive-date=2010-10-19 }}</ref><ref>{{Cite journal |last=Lehto |first=Tarja |date=1992 |title=Mycorrhizas and Drought Resistance of ''Picea sitchensis'' (Bong.) Carr. I. In Conditions of Nutrient Deficiency |journal=New Phytologist |volume=122 |issue=4 |pages=661–668 |jstor=2557434 |doi=10.1111/j.1469-8137.1992.tb00094.x |doi-access=free }}</ref><ref>{{cite journal |last1=Nikolaou |first1=N. |last2=Angelopoulos |first2=K. |last3=Karagiannidis |first3=N. |date=2003 |title=Effects of Drought Stress on Mycorrhizal and Non-Mycorrhizal Cabernet Sauvignon Grapevine, Grafted Onto Various Rootstocks |journal=Experimental Agriculture |volume=39 |issue=3 |pages=241–252 |doi=10.1017/S001447970300125X |s2cid=84997899 }}</ref> Moreover, the significance of mycorrhizal fungi also includes alleviation of salt stress and its beneficial effects on plant growth and productivity. Although salinity can negatively affect mycorrhizal fungi, many reports show improved growth and performance of mycorrhizal plants under salt stress conditions.<ref>{{cite journal |last1=Porcel |first1=Rosa |last2=Aroca |first2=Ricardo |last3=Ruiz-Lozano |first3=Juan Manuel |title=Salinity stress alleviation using arbuscular mycorrhizal fungi. A review |journal=Agronomy for Sustainable Development |date=January 2012 |volume=32 |issue=1 |pages=181–200 |doi=10.1007/s13593-011-0029-x |bibcode=2012AgSD...32..181P |s2cid=8572482 |url=https://hal.archives-ouvertes.fr/hal-00930499/file/hal-00930499.pdf }}</ref>
+[[Ecosystem services]] provided by mycorrhizal fungi may depend on the soil microbiome.<ref name="Svenningsen 1296–1307">{{cite journal |last1=Svenningsen |first1=Nanna B |last2=Watts-Williams |first2=Stephanie J |last3=Joner |first3=Erik J |last4=Battini |first4=Fabio |last5=Efthymiou |first5=Aikaterini |last6=Cruz-Paredes |first6=Carla |last7=Nybroe |first7=Ole |last8=Jakobsen |first8=Iver |title=Suppression of the activity of arbuscular mycorrhizal fungi by the soil microbiota |journal=The ISME Journal |date=May 2018 |volume=12 |issue=5 |pages=1296–1307 |doi=10.1038/s41396-018-0059-3 |pmid=29382946 |pmc=5931975 |doi-access=free |bibcode=2018ISMEJ..12.1296S }}</ref> Furthermore, mycorrhizal fungi was significantly correlated with soil physical variable, but only with water level and not with aggregate stability<ref>{{cite book |doi=10.1007/1-4020-4447-X_10 |chapter=Disease Resistance in Plants Through Mycorrhizal Fungi Induced Allelochemicals |title=Allelochemicals: Biological Control of Plant Pathogens and Diseases |series=Disease Management of Fruits and Vegetables |year=2006 |last1=Zeng |first1=Ren-Sen |volume=2 |pages=181–192 |isbn=1-4020-4445-3 }}</ref><ref>{{cite web |url=https://www.usask.ca/biology/kaminskyj/arctic.html |title=Dr. Susan Kaminskyj: Endorhizal Fungi |publisher=Usask.ca |access-date=2010-09-30 |archive-url=https://web.archive.org/web/20101104000757/http://www.usask.ca/biology/kaminskyj/arctic.html |archive-date=2010-11-04 }}</ref> and can lead also to more resistant to the effects of drought.<ref>{{cite web |url=http://aggie-horticulture.tamu.edu/Faculty/davies/research/mycorrhizae.html |title=Dr. Davies Research Page |publisher=Aggie-horticulture.tamu.edu |access-date=2010-09-30 |archive-url=https://web.archive.org/web/20101019002159/http://aggie-horticulture.tamu.edu/faculty/davies/research/mycorrhizae.html |archive-date=2010-10-19 }}</ref><ref>{{Cite journal |last=Lehto |first=Tarja |date=1992 |title=Mycorrhizas and Drought Resistance of ''Picea sitchensis'' (Bong.) Carr. I. In Conditions of Nutrient Deficiency |journal=New Phytologist |volume=122 |issue=4 |pages=661–668 |jstor=2557434 |doi=10.1111/j.1469-8137.1992.tb00094.x |doi-access=free }}</ref><ref>{{cite journal |last1=Nikolaou |first1=N. |last2=Angelopoulos |first2=K. |last3=Karagiannidis |first3=N. |date=2003 |title=Effects of Drought Stress on Mycorrhizal and Non-Mycorrhizal Cabernet Sauvignon Grapevine, Grafted Onto Various Rootstocks |journal=Experimental Agriculture |volume=39 |issue=3 |pages=241–252 |doi=10.1017/S001447970300125X |s2cid=84997899 }}</ref> Moreover, the significance of mycorrhizal fungi also includes alleviation of salt stress and its beneficial effects on plant growth and productivity. Although salinity can negatively affect mycorrhizal fungi, many reports show improved growth and performance of mycorrhizal plants under salt stress conditions.<ref>{{cite journal |last1=Porcel |first1=Rosa |last2=Aroca |first2=Ricardo |last3=Ruiz-Lozano |first3=Juan Manuel |title=Salinity stress alleviation using arbuscular mycorrhizal fungi. A review |journal=Agronomy for Sustainable Development |date=January 2012 |volume=32 |issue=1 |pages=181–200 |doi=10.1007/s13593-011-0029-x |bibcode=2012AgSD...32..181P |s2cid=8572482 |url=https://hal.archives-ouvertes.fr/hal-00930499/file/hal-00930499.pdf }}</ref>
 ===Resistance to insects===
@@ Line 164: / Line 173: @@
 == Climate change ==
-CO<sub>2</sub> released by human activities is causing [[climate change]] and possible damage to mycorrhizae, but the direct effect of an increase in the gas should be to benefit plants and mycorrhizae.<ref name="Monz Hunt Reeves Elliott 1994 pp. 75–80">{{cite journal |last1=Monz |first1=C. A. |last2=Hunt |first2=H. W. |last3=Reeves |first3=F. B. |last4=Elliott |first4=E. T. |title=The response of mycorrhizal colonization to elevated CO2 and climate change in Pascopyrum smithii and Bouteloua gracilis |journal=Plant and Soil |volume=165 |issue=1 |year=1994 |doi=10.1007/bf00009964 |pages=75–80|bibcode=1994PlSoi.165...75M |s2cid=34893610 }}</ref> In Arctic regions, nitrogen and water are harder for plants to obtain, making mycorrhizae crucial to plant growth.<ref name="Hobbie Hobbie Drossman Conte 2009 pp. 84–94">{{cite journal |last1=Hobbie |first1=John E. |last2=Hobbie |first2=Erik A. |last3=Drossman |first3=Howard |last4=Conte |first4=Maureen |last5=Weber |first5=J. C. |last6=Shamhart |first6=Julee |last7=Weinrobe |first7=Melissa |display-authors=3 |title=Mycorrhizal fungi supply nitrogen to host plants in Arctic tundra and boreal forests: 15N is the key signal|journal=Canadian Journal of Microbiology |volume=55 |issue=1 |year=2009 |doi=10.1139/w08-127 |pages=84–94|pmid=19190704 |hdl=1912/2902 |hdl-access=free }}</ref> Since mycorrhizae tend to do better in cooler temperatures, warming could be detrimental to them.<ref name="Heinemeyer Fitter 2004 pp. 525–534">{{cite journal |last1=Heinemeyer |first1=A. |last2=Fitter |first2=A. H. |title=Impact of temperature on the arbuscular mycorrhizal (AM) symbiosis: growth responses of the host plant and its AM fungal partner |journal=Journal of Experimental Botany |volume=55 |issue=396 |date=22 January 2004 |doi=10.1093/jxb/erh049 |pages=525–534|pmid=14739273 |doi-access=free }}</ref> Gases such as SO<sub>2</sub>, NO-x, and O<sub>3</sub> produced by human activity may harm mycorrhizae, causing reduction in "[[propagules]], the colonization of roots, degradation in connections between trees, reduction in the mycorrhizal incidence in trees, and reduction in the [[enzyme activity]] of ectomycorrhizal roots."<ref name="Xavier-1999">{{Cite journal |last1=Xavier |first1=L. J. |last2=Germida |first2=J. J. |title=Impact of human activities on mycorrhizae |journal=Proceedings of the 8th International Symposium on Microbial Ecology |date=1999 }}</ref>
+CO<sub>2</sub> released by human activities is causing [[climate change]] and possible damage to mycorrhizae, but the direct effect of an increase in the gas should be to benefit plants and mycorrhizae.<ref name="Monz Hunt Reeves Elliott 1994 pp. 75–80">{{cite journal |last1=Monz |first1=C. A. |last2=Hunt |first2=H. W. |last3=Reeves |first3=F. B. |last4=Elliott |first4=E. T. |title=The response of mycorrhizal colonization to elevated CO<sub>2</sub> and climate change in Pascopyrum smithii and Bouteloua gracilis |journal=Plant and Soil |volume=165 |issue=1 |year=1994 |doi=10.1007/bf00009964 |pages=75–80|bibcode=1994PlSoi.165...75M |s2cid=34893610 }}</ref> In Arctic regions, nitrogen and water are harder for plants to obtain, making mycorrhizae crucial to plant growth.<ref name="Hobbie Hobbie Drossman Conte 2009 pp. 84–94">{{cite journal |last1=Hobbie |first1=John E. |last2=Hobbie |first2=Erik A. |last3=Drossman |first3=Howard |last4=Conte |first4=Maureen |last5=Weber |first5=J. C. |last6=Shamhart |first6=Julee |last7=Weinrobe |first7=Melissa |display-authors=3 |title=Mycorrhizal fungi supply nitrogen to host plants in Arctic tundra and boreal forests: 15N is the key signal|journal=Canadian Journal of Microbiology |volume=55 |issue=1 |year=2009 |doi=10.1139/w08-127 |pages=84–94|pmid=19190704 |hdl=1912/2902 |hdl-access=free }}</ref> Since mycorrhizae tend to do better in cooler temperatures, warming could be detrimental to them.<ref name="Heinemeyer Fitter 2004 pp. 525–534">{{cite journal |last1=Heinemeyer |first1=A. |last2=Fitter |first2=A. H. |title=Impact of temperature on the arbuscular mycorrhizal (AM) symbiosis: growth responses of the host plant and its AM fungal partner |journal=Journal of Experimental Botany |volume=55 |issue=396 |date=22 January 2004 |doi=10.1093/jxb/erh049 |pages=525–534|pmid=14739273 |doi-access=free }}</ref> Gases such as SO<sub>2</sub>, NO<sub>x</sub>, and O<sub>3</sub> produced by human activity may harm mycorrhizae, causing reduction in "[[propagules]], the colonization of roots, degradation in connections between trees, reduction in the mycorrhizal incidence in trees, and reduction in the [[enzyme activity]] of ectomycorrhizal roots."<ref name="Xavier-1999">{{Cite journal |last1=Xavier |first1=L. J. |last2=Germida |first2=J. J. |title=Impact of human activities on mycorrhizae |journal=Proceedings of the 8th International Symposium on Microbial Ecology |date=1999 }}</ref>
-A company in [[Israel]], Groundwork BioAg, has discovered a method of using mycorrhizal fungi to increase agricultural crops while sequestering greenhouse gases and eliminating CO2 from the atmosphere.<ref>[https://www.haaretz.com/israel-news/2024-06-21/ty-article-magazine/.highlight/the-israeli-company-that-uses-fungus-to-tackle-the-climate-and-soil-crises/00000190-3723-d6fa-abb4-77af14b90000 he Israeli Company That Uses Fungus to Tackle the Climate and Soil Crises], ''[[Haaretz]]''</ref>
+A company in [[Israel]], Groundwork BioAg, has discovered a method of using mycorrhizal fungi to increase agricultural crops while sequestering greenhouse gases and eliminating CO<sub>2</sub> from the atmosphere.<ref>[https://www.haaretz.com/israel-news/2024-06-21/ty-article-magazine/.highlight/the-israeli-company-that-uses-fungus-to-tackle-the-climate-and-soil-crises/00000190-3723-d6fa-abb4-77af14b90000 he Israeli Company That Uses Fungus to Tackle the Climate and Soil Crises], ''[[Haaretz]]''</ref>
 == Conservation and mapping ==
-In 2021, the [[Society for the Protection of Underground Networks]] was launched. SPUN is a science-based initiative to map and protect the mycorrhizal networks regulating Earth’s climate and ecosystems. Its stated goals are mapping, protecting, and harnessing mycorrhizal fungi.
+In 2021, the [[Society for the Protection of Underground Networks]] was launched. SPUN is a science-based initiative to map and protect the mycorrhizal networks regulating Earth's climate and ecosystems. Its stated goals are mapping, protecting, and harnessing mycorrhizal fungi.
 ==See also==
@@ Line 181: / Line 190: @@
 * [[Mucigel]]
 * [[Mycorrhizal fungi and soil carbon storage]]
-* [[Mycorrhizal network]]
+* [[Mycorrhizal network|Mycorrhizal Network (Wood Wide Web)]]
 * [[Rhizobia]]
 * [[Suzanne Simard]]
@@ Line 204: / Line 213: @@
 * [http://www.mycorrhizas.org International Mycorrhiza Society] International Mycorrhiza Society
-* [http://www.ted.com/talks/mohamed_hijri_a_simple_solution_to_the_coming_phosphorus_crisis Mohamed Hijri: A simple solution to the coming phosphorus crisis] video recommending agricultural mycorrhiza use to conserve phosphorus reserves & 85% waste problem @Ted.com
+* [https://www.ted.com/talks/mohamed_hijri_a_simple_solution_to_the_coming_phosphorus_crisis Mohamed Hijri: A simple solution to the coming phosphorus crisis] video recommending agricultural mycorrhiza use to conserve phosphorus reserves & 85% waste problem @Ted.com
 * [http://mycorrhizas.info/index.html Mycorrhizal Associations: The Web Resource] Comprehensive illustrations and lists of mycorrhizal and nonmycorrhizal plants and fungi
 * [https://web.archive.org/web/20130412032458/http://www.gmo-safety.eu/science-live/439.mycorrhizas-successful-symbiosis.html Mycorrhizas – a successful symbiosis] Biosafety research into genetically modified barley

Mycorrhiza: Difference between revisions

Latest revision as of 22:23, 17 November 2025

Contents

Definition

Evolution

Emergence alongside terrestrial plants

Fossil record and genomic analysis

Origins in plants

Origins in fungi

Arbuscular mycorrhizae

Ectomycorrhizae

Orchid mycorrhizae

Ericoid mycorrhizae

Types

Similar symbiotic relationships

Ectomycorrhiza

Arbutoid mycorrhiza

Arbuscular mycorrhiza

Mucoromycotina fine root endophytes

Ericoid mycorrhiza

Orchid mycorrhiza

Monotropoid mycorrhiza

Function

Formation

Mechanisms

Sugar-water/mineral exchange

Disease, drought and salinity resistance and its correlation to mycorrhizae

Resistance to insects

Colonization of barren soil

Resistance to toxicity

Occurrence of mycorrhizal associations

Climate change

Conservation and mapping

See also

References

External links

Navigation menu

Mycorrhiza: Difference between revisions

Latest revision as of 22:23, 17 November 2025

Definition

Evolution

Emergence alongside terrestrial plants

Fossil record and genomic analysis

Origins in plants

Origins in fungi

Arbuscular mycorrhizae

Ectomycorrhizae

Orchid mycorrhizae

Ericoid mycorrhizae

Types

Similar symbiotic relationships

Ectomycorrhiza

Arbutoid mycorrhiza

Arbuscular mycorrhiza

Mucoromycotina fine root endophytes

Ericoid mycorrhiza

Orchid mycorrhiza

Monotropoid mycorrhiza

Function

Formation

Mechanisms

Sugar-water/mineral exchange

Disease, drought and salinity resistance and its correlation to mycorrhizae

Resistance to insects

Colonization of barren soil

Resistance to toxicity

Occurrence of mycorrhizal associations

Climate change

Conservation and mapping

See also

References

External links

Navigation menu

Search