Mycorrhiza

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Amanita muscaria fruit bodies.jpg
Mycorrhizal root tips (amanita).jpg
Arbuscular mycorrhiza microscope.jpg
Wheat P1210892.jpg
Many conspicuous fungi such as the fly agaric (upper left) form ectomycorrhiza (upper right) with tree rootlets. Arbuscular mycorrhiza (lower left) are very common in plants, including crop species such as wheat (lower right)

A mycorrhiza (from Greek μύκης mýkēs, "fungus", and ῥίζα rhiza, "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas [1] ) is a symbiotic association between a fungus and a plant. [2] The term mycorrhiza refers to the role of the fungus in the plant's rhizosphere, its root system. Mycorrhizae play important roles in plant nutrition, soil biology, and soil chemistry.

Contents

In a mycorrhizal association, the fungus colonizes the host plant's root tissues, either intracellularly as in arbuscular mycorrhizal fungi (AMF or AM), or extracellularly as in ectomycorrhizal fungi. [3] The association is sometimes mutualistic. In particular species or in particular circumstances, mycorrhizae may have a parasitic association with host plants. [4]

Definition

A mycorrhiza is a symbiotic association between a green plant and a fungus. The plant makes organic molecules such as sugars by photosynthesis and supplies them to the fungus, and the fungus supplies to the plant 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 [5] and there is fossil evidence that early land plants that lacked roots formed arbuscular mycorrhizal associations. [6] 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 wheat and rice. [7]

Evolution

Fossil and genetic evidence indicate that mycorrhizae are ancient, potentially as old as the terrestrialization of plants. Genetic evidence indicates that all land plants share a single common ancestor, [8] which appears to have quickly adopted mycorrhizal symbiosis, and research suggests that proto-mycorrhizal fungi were a key factor enabling plant terrestrialization. [9] 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 major, giving a lower bound for how late mycorrhizal symbiosis may have developed. [6] Ectomycorrhizae developed substantially later, during the Jurassic period, while most other modern mycorrhizal families, including orchid and erchoid mycorrhizae, date to the period of angiosperm radiation in the Cretaceous period. [10] There is genetic evidence that the symbiosis between legumes and nitrogen-fixing bacteria is an extension of mycorrhizal symbiosis. [11] 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. [12]

Types

Mycorrhizas are commonly divided into ectomycorrhizas and endomycorrhizas. The two types are differentiated by the fact that the hyphae of ectomycorrhizal fungi do not penetrate individual cells within the root, while the hyphae of endomycorrhizal fungi penetrate the cell wall and invaginate the cell membrane. [13] [14] Endomycorrhiza includes arbuscular, ericoid, and orchid mycorrhiza, while arbutoid mycorrhizas can be classified as ectoendomycorrhizas. Monotropoid mycorrhizas form a special category.

Ectomycorrhiza

Beech is ectomycorrhizal Grib skov.jpg
Beech is ectomycorrhizal
Leccinum aurantiacum, an ectomycorrhizal fungus Raudonvirsis1-vi.jpg
Leccinum aurantiacum , an ectomycorrhizal fungus

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 [15] families, orchids, [16] and fungi belonging to the Basidiomycota, Ascomycota, and Zygomycota. Some EcM fungi, such as many Leccinum and Suillus , are symbiotic with only one particular genus of plant, while other fungi, such as the Amanita , are generalists that form mycorrhizas with many different plants. [17] An individual tree may have 15 or more different fungal EcM partners at one time. [18] 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. [19]

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

Nutrients can be shown to move between different plants through the fungal network. Carbon has been shown to move from paper birch trees into Douglas-fir trees thereby promoting succession in ecosystems. [20] 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. [21] [22] When compared with non-mycorrhizal fine roots, ectomycorrhizae may contain very high concentrations of trace elements, including toxic metals (cadmium, silver) or chlorine. [23]

The first genomic sequence for a representative of symbiotic fungi, the ectomycorrhizal basidiomycete L. bicolor, was published in 2008. [24] 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.

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.[ citation needed ] It differs from ectomycorrhiza in that some hyphae actually penetrate into the root cells, making this type of mycorrhiza an ectendomycorrhiza. [25]

Endomycorrhiza

Endomycorrhizas are variable and have been further classified as arbuscular, ericoid, arbutoid, monotropoid, and orchid mycorrhizas. [26]

Wheat is arbuscular mycorrhizal Wheat field.jpg
Wheat is arbuscular mycorrhizal

Arbuscular mycorrhiza

Arbuscular mycorrhizas, or AM (formerly known as vesicular-arbuscular mycorrhizas, or VAM), are mycorrhizas whose hyphae penetrate plant cells, producing structures that are either balloon-like (vesicles) or dichotomously branching invaginations (arbuscules) as a means of nutrient exchange. The fungal hyphae do not in fact penetrate the protoplast (i.e. the interior of the cell), but invaginate the cell membrane. The structure of the arbuscules greatly increases the contact surface area between the hypha and the cell cytoplasm to facilitate the transfer of nutrients between them.

Arbuscular mycorrhizas are formed only by fungi in the division Glomeromycota. Fossil evidence [6] and DNA sequence analysis [27] 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. [15] The hyphae of arbuscular mycorrhizal fungi produce the glycoprotein glomalin, which may be one of the major stores of carbon in the soil. [28] Arbuscular mycorrhizal fungi have (possibly) been asexual for many millions of years and, unusually, individuals can contain many genetically different nuclei (a phenomenon called heterokaryosis). [29]

Ericoid mycorrhiza

An ericoid mycorrhizal fungus isolated from Woollsia pungens Ericoid mycorrhizal fungus.jpg
An ericoid mycorrhizal fungus isolated from Woollsia pungens

Ericoid mycorrhizas are the third of the three more ecologically important types. They 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. [14]

Ericoid mycorrhizas have also been shown to have considerable saprotrophic capabilities, which would enable plants to receive nutrients from not-yet-decomposed materials via the decomposing actions of their ericoid partners. [31]

Orchid mycorrhiza

All orchids are myco-heterotrophic at some stage during their lifecycle and form orchid mycorrhizas with a range of basidiomycete fungi.[ citation needed ] Their hyphae penetrate into the root cells and form pelotons (coils) for nutrient exchange.[ citation needed ]

Monotropoid mycorrhiza

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.[ citation needed ]

Mutualist dynamics

Nutrient exchanges and communication between a mycorrhizal fungus and plants. 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. [32]

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. [33] It has also been suggested that evolutionary and phylogenetic relationships can explain much more variation in the strength of mycorrhizal mutualisms than ecological factors. [34]

Within mutualistic mycorrhiza, the plant gives carbohydrates (products of photosynthesis) to the fungus, while the fungus gives the plant water and minerals in exchange. Mutualistic mycorrhiza en.svg
Within mutualistic mycorrhiza, the plant gives carbohydrates (products of photosynthesis) to the fungus, while the fungus gives the plant water and minerals in exchange.

Sugar-water/mineral exchange

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).</ref> 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).</ref>

The mycorrhizal mutualistic association provides the fungus with relatively constant and direct access to carbohydrates, such as glucose and sucrose. [35] 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. [36]

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. [37] 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. [38] Inga alley cropping , proposed as an alternative to slash and burn rainforest destruction, [39] relies upon mycorrhiza within the root system of species of Inga to prevent the rain from washing phosphorus out of the soil. [40]

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. [41]

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. [42]

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. [43] Mycorrhizae are especially beneficial for the plant partner in nutrient-poor soils. [44]

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. [45] 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. [46]

AMF was also significantly correlated with soil biological fertility variables such as soil microbial communities and associated disease suppressiveness. [47] Thus, ecosystem services provided by AMF may depend on the soil microbiome. [47] Furthermore, AMF was significantly correlated with soil physical variable, but only with water level and not with aggregate stability. [48] [49] and are also more resistant to the effects of drought. [50] [51] [52] The significance of arbuscular mycorrhizal fungi includes alleviation of salt stress and its beneficial effects on plant growth and productivity. Although salinity can negatively affect arbuscular mycorrhizal fungi, many reports show improved growth and performance of mycorrhizal plants under salt stress conditions. [53]

Resistance to insects

Research has shown that plants connected by mycorrhizal fungi can use these underground connections to produce and receive warning signals. [54] [55] Specifically, when a host plant is attacked by an aphid, the plant signals surrounding connected plants of its condition. The host plant releases volatile organic compounds (VOCs) that attract the insect's predators. The plants connected by mycorrhizal fungi are also prompted to produce identical VOCs that protect the uninfected plants from being targeted by the insect. [54] Additionally, this assists the mycorrhizal fungi by preventing the plant's carbon relocation which negatively affects the fungi's growth and occurs when the plant is attacked by herbivores. [54]

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. [56] The absence of mycorrhizal fungi can also slow plant growth in early succession or on degraded landscapes. [57] The introduction of alien mycorrhizal plants to nutrient-deficient ecosystems puts indigenous non-mycorrhizal plants at a competitive disadvantage. [58] This aptitude to colonize barren soil is defined by the category Oligotroph.

Resistance to toxicity

Fungi have been found to have a protective role for plants rooted in soils with high metal concentrations, such as acidic and contaminated soils. Pine trees inoculated with Pisolithus tinctorius planted in several contaminated sites displayed high tolerance to the prevailing contaminant, survivorship and growth. [59] 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. [58]

Climate change

This section is an excerpt from Mycorrhizae and climate change.[ edit ]

Mycorrhizae and climate change refers to the effects of climate change on mycorrhizae, a fungus which forms an endosymbiotic relationship between with a vascular host plant [60] by colonizing its roots, and the effects brought on by climate change. Climate change is any lasting effect in weather or temperature. It is important to note that a good indicator of climate change is global warming, though the two are not analogous. [61] However, temperature plays a very important role in all ecosystems on Earth, especially those with high counts of mycorrhiza in soil biota.

Mycorrhizae are one of the most widespread symbioses on the planet, as they form a plant-fungal interaction with nearly eighty percent of all terrestrial plants. [62] The resident mycorrhizae benefits from a share of the sugars and carbon produced during photosynthesis, while the plant effectively accesses water and other nutrients, such as nitrogen and phosphorus, crucial to its health. [63] This symbiosis has become so beneficial to terrestrial plants that some depend entirely on the relationship to sustain themselves in their respective environments. The fungi are essential to the planet as most ecosystems, especially those in the Arctic, are filled with plants that survive with the aid of mycorrhizae. Because of their importance to a productive ecosystem, understanding this fungus and its symbioses is currently an active area of scientific research.

Occurrence of mycorrhizal associations

Mycorrhizas are present in 92% of plant families studied (80% of species), [15] with arbuscular mycorrhizas being the ancestral and predominant form, [15] and the most prevalent symbiotic association found in the plant kingdom. [35] The structure of arbuscular mycorrhizas has been highly conserved since their first appearance in the fossil record, [6] with both the development of ectomycorrhizas, and the loss of mycorrhizas, evolving convergently on multiple occasions. [15]

Discovery

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. [64] This symbiosis was studied and described by Franciszek Kamieński in 1879–1882. [65] [66]

See also

Related Research Articles

<span class="mw-page-title-main">Arbuscular mycorrhiza</span> Symbiotic penetrative association between a fungus and the roots of a vascular plant

An arbuscular mycorrhiza (AM) is a type of mycorrhiza in which the symbiont fungus penetrates the cortical cells of the roots of a vascular plant forming arbuscules.

<span class="mw-page-title-main">Root hair</span> Part of plant root

Root hair, or absorbent hairs, are outgrowths of epidermal cells, specialized cells at the tip of a plant root. They are lateral extensions of a single cell and are only rarely branched. They are found in the region of maturation, of the root. Root hair cells improve plant water absorption by increasing root surface area to volume ratio which allows the root hair cell to take in more water. The large vacuole inside root hair cells makes this intake much more efficient. Root hairs are also important for nutrient uptake as they are main interface between plants and mycorrhizal fungi.

<span class="mw-page-title-main">Glomeromycota</span> Phylum of fungi

Glomeromycota are one of eight currently recognized divisions within the kingdom Fungi, with approximately 230 described species. Members of the Glomeromycota form arbuscular mycorrhizas (AMs) with the thalli of bryophytes and the roots of vascular land plants. Not all species have been shown to form AMs, and one, Geosiphon pyriformis, is known not to do so. Instead, it forms an endocytobiotic association with Nostoc cyanobacteria. The majority of evidence shows that the Glomeromycota are dependent on land plants for carbon and energy, but there is recent circumstantial evidence that some species may be able to lead an independent existence. The arbuscular mycorrhizal species are terrestrial and widely distributed in soils worldwide where they form symbioses with the roots of the majority of plant species (>80%). They can also be found in wetlands, including salt-marshes, and associated with epiphytic plants.

Glomus is a genus of arbuscular mycorrhizal (AM) fungi, and all species form symbiotic relationships (mycorrhizae) with plant roots. Glomus is the largest genus of AM fungi, with ca. 85 species described, but is currently defined as non-monophyletic.

<span class="mw-page-title-main">Ericoid mycorrhiza</span> Species of fungus

The ericoid mycorrhiza is a mutualistic relationship formed between members of the plant family Ericaceae and several lineages of mycorrhizal fungi. This symbiosis represents an important adaptation to acidic and nutrient poor soils that species in the Ericaceae typically inhabit, including boreal forests, bogs, and heathlands. Molecular clock estimates suggest that the symbiosis originated approximately 140 million years ago.

Microbial inoculants also known as soil inoculants or bioinoculants are agricultural amendments that use beneficial rhizosphericic or endophytic microbes to promote plant health. Many of the microbes involved form symbiotic relationships with the target crops where both parties benefit (mutualism). While microbial inoculants are applied to improve plant nutrition, they can also be used to promote plant growth by stimulating plant hormone production. Although bacterial and fungal inoculants are common, inoculation with archaea to promote plant growth is being increasingly studied.

<span class="mw-page-title-main">Hartig net</span> Network of inward-growing hyphae

Hartig net is a network of inward-growing hyphae, that extends into the root, penetrating between the epidermis and cortex of ectomycorrhizal plants. This network is a site of nutrient exchange between the fungus and the host plant. The Hartig net is one of the three components required for ectomycorrhizal roots to form as part of ectomycorrhizal symbiosis with the host tree or plant.

Nitrogen nutrition in the arbuscular mycorrhizal system refers to...

The mycorrhizosphere is the region around a mycorrhizal fungus in which nutrients released from the fungus increase the microbial population and its activities. The roots of most terrestrial plants, including most crop plants and almost all woody plants, are colonized by mycorrhiza-forming symbiotic fungi. In this relationship, the plant roots are infected by a fungus, but the rest of the fungal mycelium continues to grow through the soil, digesting and absorbing nutrients and water and sharing these with its plant host. The fungus in turn benefits by receiving photosynthetic sugars from its host. The mycorrhizosphere consists of roots, hyphae of the directly connected mycorrhizal fungi, associated microorganisms, and the soil in their direct influence.

<span class="mw-page-title-main">Mycorrhizal network</span> Underground hyphal networks that connect individual plants together

A Mycorrhizal network is an underground network found in forests and other plant communities, created by the hyphae of mycorrhizal fungi joining with plant roots. This network connects individual plants together and transfers water, carbon, nitrogen, and other nutrients and minerals between participants. Several studies have demonstrated that mycorrhizal networks can transport carbon, phosphorus, nitrogen, water, defense compounds, and allelochemicals from plant to plant. The flux of nutrients and water through hyphal networks has been proposed to be driven by a source–sink model, where plants growing under conditions of relatively high resource availability transfer carbon or nutrients to plants located in less favorable conditions. A common example is the transfer of carbon from plants with leaves located in high-light conditions in the forest canopy, to plants located in the shaded understory where light availability limits photosynthesis. In natural ecosystems, plants may be dependent on fungal symbionts for 90% of their phosphorus requirements and 80% of their nitrogen requirements. Mycorrhizal relationships are most commonly mutualistic, with both partners benefiting, but can be commensal or parasitic.

<span class="mw-page-title-main">Mycorrhizal fungi and soil carbon storage</span>

Soil carbon storage is an important function of terrestrial ecosystems. Soil contains more carbon than plants and the atmosphere combined. Understanding what maintains the soil carbon pool is important to understand the current distribution of carbon on Earth, and how it will respond to environmental change. While much research has been done on how plants, free-living microbial decomposers, and soil minerals affect this pool of carbon, it is recently coming to light that mycorrhizal fungi—symbiotic fungi that associate with roots of almost all living plants—may play an important role in maintaining this pool as well. Measurements of plant carbon allocation to mycorrhizal fungi have been estimated to be 5 to 20% of total plant carbon uptake, and in some ecosystems the biomass of mycorrhizal fungi can be comparable to the biomass of fine roots. Recent research has shown that mycorrhizal fungi hold 50 to 70 percent of the total carbon stored in leaf litter and soil on forested islands in Sweden. Turnover of mycorrhizal biomass into the soil carbon pool is thought to be rapid and has been shown in some ecosystems to be the dominant pathway by which living carbon enters the soil carbon pool.

<span class="mw-page-title-main">Ectomycorrhiza</span> Non-penetrative symbiotic association between a fungus and the roots of a vascular plant

An ectomycorrhiza is a form of symbiotic relationship that occurs between a fungal symbiont, or mycobiont, and the roots of various plant species. The mycobiont is often from the phyla Basidiomycota and Ascomycota, and more rarely from the Zygomycota. Ectomycorrhizas form on the roots of around 2% of plant species, usually woody plants, including species from the birch, dipterocarp, myrtle, beech, willow, pine and rose families. Research on ectomycorrhizas is increasingly important in areas such as ecosystem management and restoration, forestry and agriculture.

<i>Rhizophagus irregularis</i> Species of arbuscular mycorrhizal fungus used as a soil inoculant in agriculture and horticulture

Rhizophagus irregularis is an arbuscular mycorrhizal fungus used as a soil inoculant in agriculture and horticulture. Rhizophagus irregularis is also commonly used in scientific studies of the effects of arbuscular mycorrhizal fungi on plant and soil improvement. Until 2001, the species was known and widely marketed as Glomus intraradices, but molecular analysis of ribosomal DNA led to the reclassification of all arbuscular fungi from Zygomycota phylum to the Glomeromycota phylum.

<span class="mw-page-title-main">Ectomycorrhizal extramatrical mycelium</span>

Ectomycorrhizal extramatrical mycelium is the collection of filamentous fungal hyphae emanating from ectomycorrhizas. It may be composed of fine, hydrophilic hypha which branches frequently to explore and exploit the soil matrix or may aggregate to form rhizomorphs; highly differentiated, hydrophobic, enduring, transport structures.

<span class="mw-page-title-main">Root microbiome</span>

The root microbiome is the dynamic community of microorganisms associated with plant roots. Because they are rich in a variety of carbon compounds, plant roots provide unique environments for a diverse assemblage of soil microorganisms, including bacteria, fungi and archaea. The microbial communities inside the root and in the rhizosphere are distinct from each other, and from the microbial communities of bulk soil, although there is some overlap in species composition.

<i>Cenococcum geophilum</i> Species of fungus

Cenococcum geophilum Fr., synonym Cenococcum graniforme (Sow.) Ferd. and Winge, is an Ascomycete fungal species and is the only member in the genus Cenococcum. It is one of the most common ectomycorrhizal fungal species encountered in forest ecosystems. The geographic distribution of the species is notably cosmopolitan; it is found in ecosystems with a wide range of environmental conditions, and in many cases in high relative frequency. Because of its wide distribution and abundance in forest soils, it is one of the most well-studied ectomycorrhizal fungal species. While the species has long been known to be sterile and not produce asexual or sexual spores, cryptic sexual stages may exist. The hyphae produced by C. geophilum are characterized by their thick (1.5-8 um), straight and jet black appearance with little branching. They usually form monopodial (unbranched) ectomycorrhizas. The mantles of C. geophilum ectomycorrhizas are usually thick with few to many emanating hyphae.

Dark septate endophytes (DSE) are a group of endophytic fungi characterized by their morphology of melanized, septate, hyphae. This group is likely paraphyletic, and contain conidial as well as sterile fungi that colonize roots intracellularly or intercellularly. Very little is known about the number of fungal taxa within this group, but all are in the Ascomycota. They are found in over 600 plant species and across 114 families of angiosperms and gymnosperms and co-occur with other types of mycorrhizal fungi. They have a wide global distribution and can be more abundant in stressed environments. Much of their taxonomy, physiology, and ecology are unknown.

Orchid mycorrhizae are endomycorrhizal fungi which develop symbiotic relationships with the roots and seeds of plants of the family Orchidaceae. Nearly all orchids are myco-heterotrophic at some point in their life cycle. Orchid mycorrhizae are critically important during orchid germination, as an orchid seed has virtually no energy reserve and obtains its carbon from the fungal symbiont.

<span class="mw-page-title-main">Mycorrhiza helper bacteria</span> Group of organisms

Mycorrhiza helper bacteria (MHB) are a group of organisms that form symbiotic associations with both ectomycorrhiza and arbuscular mycorrhiza. MHBs are diverse and belong to a wide variety of bacterial phyla including both Gram-negative and Gram-positive bacteria. Some of the most common MHBs observed in studies belong to the phylas Pseudomonas and Streptomyces. MHBs have been seen to have extremely specific interactions with their fungal hosts at times, but this specificity is lost with plants. MHBs enhance mycorrhizal function, growth, nutrient uptake to the fungus and plant, improve soil conductance, aid against certain pathogens, and help promote defense mechanisms. These bacteria are naturally present in the soil, and form these complex interactions with fungi as plant root development starts to take shape. The mechanisms through which these interactions take shape are not well-understood and needs further study.

Mycorrhizae and climate change refers to the effects of climate change on mycorrhizae, a fungus which forms an endosymbiotic relationship between with a vascular host plant by colonizing its roots, and the effects brought on by climate change. Climate change is any lasting effect in weather or temperature. It is important to note that a good indicator of climate change is global warming, though the two are not analogous. However, temperature plays a very important role in all ecosystems on Earth, especially those with high counts of mycorrhiza in soil biota.

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