Glomeromycota

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Glomeromycota
Gigaspora margarita.JPG
Gigaspora margarita in association with Lotus corniculatus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Glomeromycota
Subdivision: Glomeromycotina
C.Walker & A.Schuessler (2001) [1]
Class: Glomeromycetes
Caval.-Sm. (1998) [2]
Orders

Glomeromycota (often referred to as glomeromycetes, as they include only one class, Glomeromycetes) are one of eight currently recognized divisions within the kingdom Fungi, [3] with approximately 230 described species. [4] 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. [5] The majority of evidence shows that the Glomeromycota are dependent on land plants (Nostoc in the case of Geosiphon) for carbon and energy, but there is recent circumstantial evidence that some species may be able to lead an independent existence. [6] 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.

Contents

According to multigene phylogenetic analyses, this taxon is located as a member of the phylum Mucoromycota. [7] Currently, the phylum name Glomeromycota is invalid, and the subphylum Glomeromycotina should be used to describe this taxon. [8]

Reproduction

The Glomeromycota have generally coenocytic (occasionally sparsely septate) mycelia and reproduce asexually through blastic development of the hyphal tip to produce spores [1] (Glomerospores) with diameters of 80–500  μm. [9] In some, complex spores form within a terminal saccule. [1] Recently it was shown that Glomus species contain 51 genes encoding all the tools necessary for meiosis. [10] Based on these and related findings, it was suggested that Glomus species may have a cryptic sexual cycle. [10] [11] [12]

Colonization

New colonization of AM fungi largely depends on the amount of inoculum present in the soil. [13] Although pre-existing hyphae and infected root fragments have been shown to successfully colonize the roots of a host, germinating spores are considered to be the key players in new host establishment. Spores are commonly dispersed by fungal and plant burrowing herbivore partners, but some air dispersal capabilities are also known. [14] Studies have shown that spore germination is specific to particular environmental conditions such as right amount of nutrients, temperature or host availability. It has also been observed that the rate of root system colonization is directly correlated to spore density in the soil. [13] In addition, new data also suggests that AM fungi host plants also secrete chemical factors which attract and enhance the growth of developing spore hyphae towards the root system. [14]

The necessary components for the colonization of Glomeromycota include, the host's fine root system, proper development of intracellular arbuscular structures, and a well-established external fungal mycelium. Colonization is accomplished by the interactions between germinating spore hyphae and the root hairs of the host or by development of appressoria between epidermal root cells. The process is regulated by specialized chemical signaling and by changes in gene expression of both the host and AM fungi. Intracellular hyphae extend up to the cortical cells of the root and penetrate the cell walls, but not the inner cellular membrane creating an internal invagination. The penetrating hyphae develop a highly branched structure called an arbuscule which have low functional periods before degradation and absorption by host's root cells. A fully developed arbuscular mycorrhizal structure facilitates the two-way movement of nutrients between the host and mutualistic fungal partner. The symbiotic association allows the host plant to respond better to environment stresses, and the non-photosynthetic fungi to obtain carbohydrates produced by photosynthesis. [14]

Phylogeny

Initial studies of the Glomeromycota were based on the morphology of soil-borne sporocarps (spore clusters) found in or near colonized plant roots. [15] Distinguishing features such as wall morphologies, size, shape, color, hyphal attachment and reaction to staining compounds allowed a phylogeny to be constructed. [16] Superficial similarities led to the initial placement of genus Glomus in the unrelated family Endogonaceae. [17] Following broader reviews that cleared up the sporocarp confusion, the Glomeromycota were first proposed in the genera Acaulospora and Gigaspora [18] before being accorded their own order with the three families Glomaceae (now Glomeraceae), Acaulosporaceae and Gigasporaceae. [19]

With the advent of molecular techniques this classification has undergone major revision. An analysis of small subunit (SSU) rRNA sequences [20] indicated that they share a common ancestor with the Dikarya. [1] Nowadays it is accepted that Glomeromycota consists of 4 orders. [21]

Glomeromycota

  Diversisporales

  Glomerales

  Archaeosporales

  Paraglomerales

Several species which produce glomoid spores (i.e. spores similar to Glomus ) in fact belong to other deeply divergent lineages [22] and were placed in the orders, Paraglomerales and Archaeosporales. [1] This new classification includes the Geosiphonaceae, which presently contains one fungus (Geosiphon pyriformis) that forms endosymbiotic associations with the cyanobacterium Nostoc punctiforme [23] and produces spores typical to this division, in the Archaeosporales.

Work in this field is incomplete, and members of Glomus may be better suited to different genera [24] or families. [9]


Molecular biology

The biochemical and genetic characterization of the Glomeromycota has been hindered by their biotrophic nature, which impedes laboratory culturing. This obstacle was eventually surpassed with the use of root cultures and, most recently, a method which applies sequencing of single nucleus from spores has also been developed to circumvent this challenge. [25] The first mycorrhizal gene to be sequenced was the small-subunit ribosomal RNA (SSU rRNA). [26] This gene is highly conserved and commonly used in phylogenetic studies so was isolated from spores of each taxonomic group before amplification through the polymerase chain reaction (PCR). [27] A metatranscriptomic survey of the Sevilleta Arid Lands found that 5.4% of the fungal rRNA reads mapped to Glomeromycota. This result was inconsistent with previous PCR-based studies of community structure in the region, suggesting that previous PCR-based studies may have underestimated Glomeromycota abundance due to amplification biases. [28]

See also

Related Research Articles

<span class="mw-page-title-main">Mycorrhiza</span> Fungus-plant symbiotic association

A mycorrhiza is a symbiotic association between a fungus and a plant. 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.

<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. Arbuscular mycorrhiza is a type of endomycorrhiza along with ericoid mycorrhiza and orchid mycorrhiza .They are characterized by the formation of unique tree-like structures, the arbuscules. In addition, globular storage structures called vesicles are often encountered.

Glomus aggregatum is an arbuscular mycorrhizal fungus used as a soil inoculant in agriculture and horticulture. Like other species in this phylum it forms obligate symbioses with plant roots, where it obtains carbon (photosynthate) from the host plant in exchange for nutrients and other benefits.

<i>Glomus</i> (fungus) Genus of arbuscular mycorrhizal fungi

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.

<i>Geosiphon</i> Monotypic genus of photosynthetic, non-lichen fungi

Geosiphon is a genus of fungus in the family Geosiphonaceae. The genus is monotypic, containing the single species Geosiphon pyriformis, first described by Kützing in 1849 as Botrydium pyriforme. In 1915, Von Wettstein characterized Geosiphon pyriforme as a multinucleate alga containing endosymbiotic cyanobacteria, although he also noted the presence of chitin, a component of fungal cell walls. In 1933, Knapp was the first to suggest the fungal origin of the species and described it as a lichen with endosymbiotic cyanobacteria. It is the only member of the Glomeromycota known to not form a symbiosis with terrestrial plants in the form of arbuscular mycorrhiza.

Archaeosporales is an order of fungi best known as arbuscular mycorrhiza to vascular land plants (Tracheophyta). But also form free living endocyte symbioses with cyanobacteria. The free living forms have a Precambrian fossil record back 2.2 Ga, well before evolution of Tracheophyta.

The Paraglomerales are a group of exclusively hypogeous (underground) arbuscular mycorrhizal fungi that rarely produce vesicles and reproduce through unpigmented spores. It includes the species Paraglomus brasilianum, Paraglomus laccatum, and Paraglomus occultum.

<span class="mw-page-title-main">Glomerales</span> Order of fungi

Glomerales is an order of symbiotic fungi within the phylum Glomeromycota.

<span class="mw-page-title-main">Mucoromycotina</span> Subphylum of fungi

Mucoromycotina is a subphylum of uncertain placement in Fungi. It was considered part of the phylum Zygomycota, but recent phylogenetic studies have shown that it was polyphyletic and thus split into several groups, it is now thought to be a paraphyletic grouping. Mucoromycotina is currently composed of 3 orders, 61 genera, and 325 species. Some common characteristics seen throughout the species include: development of coenocytic mycelium, saprotrophic lifestyles, and filamentous.

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">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.

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.

<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.

<span class="mw-page-title-main">Mucoromycota</span> Diverse group of molds

Mucoromycota is a division within the kingdom fungi. It includes a diverse group of various molds, including the common bread molds Mucor and Rhizopus. It is a sister phylum to Dikarya.

<i>Funneliformis mosseae</i> Species of fungus

Funneliformis mosseae is a species of fungus in the family Glomeraceae, which is an arbuscular mycorrhizal (AM) fungi that forms symbiotic relationships with plant roots. Funneliformis mosseae has a wide distribution worldwide, and can be found in North America, South America, Europe, Africa, Asia and Australia. Funneliformis are characterized by having an easily visible septum in the area of the spore base and are often cylindrical or funnel-shaped. Funneliformis mosseae similarly resembles Glomus caledonium, however the spore wall of Funneliformis mosseae contains three layers, whereas Gl. caledonium spore walls are composed of four layers. Funneliformis is an easily cultivated species which multiplies well in trap culture, along with its high distribution, F. mosseae is not considered endangered and is often used for experimental purposes when combined with another host.

Rhizophagus clarus is an arbuscular mycorrhizal fungus in the family Glomeraceae. The species has been shown to improve nutrient absorption and growth in several agricultural crops but is not typically applied commercially.

The International Collection of (Vesicular) Arbuscular Mycorrhizal Fungi (INVAM) is the largest collection of living arbuscular mycorrhizal fungi (AMF) and includes Glomeromycotan species from 6 continents. Curators of INVAM acquire, grow, identify, and elucidate the biology, taxonomy, and ecology of a diversity AMF with the mission to expand availability and knowledge of these symbiotic fungi. Culturing AMF presents difficulty as these fungi are obligate biotrophs that must complete their life cycle while in association with their plant hosts, while resting spores outside of the host are vulnerable to predation and degradation. Curators of INVAM have thus developed methods to overcome these challenges to increase the availability of AMF spores. The inception of this living collection of germplasm occurred in the 1980s and it takes the form of fungi growing in association with plant symbionts in the greenhouse, with spores preserved in cold storage within their associated rhizosphere. AMF spores acquired from INVAM have been used extensively in both basic and applied research projects in the fields of ecology, evolutionary biology, agroecology, and in restoration. INVAM is umbrellaed under the Kansas Biological Survey at The University of Kansas, an R1 Research Institution. The Kansas Biological Survey is also home to the well-known organization Monarch Watch. INVAM is currently located within the tallgrass prairie ecoregion, and many collaborators and researchers associated with INVAM study the role of AMF in the mediation of prairie biodiversity. James Bever and Peggy Schultz are the Curator and Director of Operation team, with Elizabeth Koziol and Terra Lubin as Associate Curators.

Ambispora granatensis is an arbuscular mycorrhizal fungal species in the genus Ambispora, family Ambisporaceae. It forms spores of the acaulosporois and glomoid morphs, thus the Ambispora classification. It was discovered in Granada Spain in 2010 and has unique spore characteristics, which distinguishes the species from the others in its genus.

<i>Glomus macrocarpum</i> Species of fungus

Glomus macrocarpum is a vesicular-arbuscular endomycorrhizal plant pathogen in the Glomeraceae family of fungi. Also occasionally known as Endogone macrocarpa, G. macrocarpum is pathogenic to multiple plants, including tobacco and chili plants. G. macrocarpum was first discovered in the French woodlands by the Tulsane brothers in the early to mid 1800s. Their first known description of G. macrocarpum was published in the New Italian Botanical Journal in 1845. G. macrocarpum has since been documented in over 26 countries, including Australia, China, and Japan for example. G. macrocarpum is frequently found in grassy meadows, forests, greenhouses, and fruit orchards. It is known for its small, round-edged, and light brown to yellow-brown sporocarp. G. macrocarpum is sometimes known as the Glomerales truffle.

References

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