Gigaspora margarita | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Fungi |
Division: | Glomeromycota |
Class: | Glomeromycetes |
Order: | Diversisporales |
Family: | Gigasporaceae |
Genus: | Gigaspora |
Species: | G. margarita |
Binomial name | |
Gigaspora margarita (W.N.) Becker & (I.R.) Hall 1976 | |
Gigaspora margarita is an Arbuscular Mycorrhizal Fungi (AMF) which means it is an obligate symbiont that creates mutualistic relationships with many different plant species. Being an AMF, G. margarita does not produce a fruiting body. All of its mycelium will be found in the soil, associating with plant roots. Though hard to distinguish between different species of AMF, microscopic distinctions can be made. A prominent morphological distinction for species in the Gigasporaceae family is their large sized spores. Gigaspora margarita is characterized by its large, white, pearl-like spores found anywhere from 260 - 400 micrometers. [1] This is where it gets its name as margarita in Latin means pearl.
Associating with many plants, Gigaspora margarita has been found in diverse regions across the globe. In culture, G. margarita has been found to associate with onion, tomato, soy beans, corn, and clover although this list is probably a lot longer. [2] Furthermore, G. margarita also associates with endobacteria making it a metaorganism that serves as a connection of three different kingdoms (plant, bacteria, fungus). Strains of G. margarita isolated without the endobacteria are possible indicating an asymmetric association between the fungi and the endobacteria. However these, ‘cured’ strains of G. margarita do not interact with its associated plants as well as strains with the endobacteria. [3]
Arbuscular Mycorrhizal Fungi can be hard to distinguish since they do not produce fruiting bodies and their entire lifecycle is completed below ground. Researchers distinguish species by looking at their microscopic morphologies and genetics.
Arbuscular Mycorrhizal Fungi are characterized by their intracellular arbuscules they form within the associated plant's roots. These arbuscules can come in many different variations with no one looking identical to another. This makes it hard for them to be used as a classification tool. Arbuscules are bush-like structures where they have branches hyphae forming from a swollen hyphal base. Oftentimes, the hyphae of AMF will stain blue using trypan blue dye. [1]
Gigaspora margarita is distinguished primarily by the morphology of its spores. Young spores are often salmon colored and will become pearly white to yellow-brown at maturity. A mature spore has three cell wall layers (L1, L2, L3):
Gigaspora margarita also has auxiliary cells produced on tightly wound hyphae. These cells are spikey in appearance and are found in clusters of 4-20. Auxiliary cells are found in all species classified under the Gigasporaceae family so while they do not specifically distinguish G. margarita, they are a good indicator that a certain AMF species is in the Gigasporaceae family. [4]
Gigaspora margarita has a peculiar genetic makeup as its genome consist of around 831 Mega base pairs (Mbp). [5] This is massive compared to the usual fungal genome size that ranges from 8.97 Mb to 177.57 Mb. [6] Furthermore, the 10 largest genomes in the kingdom of fungi belong to species that are either obligate biotrophs, endophytes, or gut fungi. This may seem like an indication that large genomes correlate with better symbiotic relationships with plants however this is not necessarily the case. The composition of the genome of G. margarita is also unique as it is primarily made up of transposable elements (64%). Fungi usually have low levels of transposable elements often only making up 0-25% of the genome. It seems the only other fungi to have large transposable element concentrations are plant pathogens which makes sense because they allow the species to adapt quickly often as a means of overcoming plant defenses. For an AMF fungus, the reasons for having such a large repertoire of transposable elements remains unclear. The genome was also found to consist of gene sequences called Helitrons. Their purpose remains unknown however these indicate that G. margarita may have captured genes from other organisms at some point in time. [5]
Gigaspora margarita has a unique ability to associate with a diverse range of endobacteria. [7] Though rare, the ability for Gigaspora margarita to host this intrahyphal bacteria is not completely unique. Many forms of AMF have been shown to associate with bacteria. The primary species of bacteria that associates with G. margarita is named Candidatus Glomeribacter gigasporarum (italicized?). Found in the Burkholderia genus, Ca G gigasporarum has had its entire genome sequenced and found to be a lot smaller than other Burkholderia species. [8] This indicates a reliance on G. sporangium to survive. Ca G gigasporarum has never been found on its own in nature and, so far, can not be isolated in culture. Adding to this hypothesis is the fact that, Ca. G. gigasporarum is vertically transmitted with G. margarita spores. Each spore contains around 250,000 of these bacteria which may be a contributor to the unusually large spores G. margarita produces. [9]
The symbiotic relationship between G. margarita and its endobacteria is asymmetric meaning one of the species can survive without the other while the other species cannot. In this case, G. margarita has been isolated and grown without the presence of endobacteria. Still, endobacteria play an important role in optimizing the relationship between the plant and the fungus. The presence of endobacteria is correlated with a higher antioxidant metabolic rate and lipid biosynthesis in the associated plant. The synthesized lipids would end up being exchanged to G. margarita. [3] Basically, endobacteria are not required for the completion of the G. margarita life cycle however they are major contributors to the health of the interaction between plant and fungus.
Little is known about the habitat and distribution other than it seems to be widespread. G. margarita is found across the globe and has been confirmed in Brazil, USA, Canada, China, Cuba, India, Japan, Mexico, New Zealand, Poland, Syria. Most extensive research on its habitat has been conducted in Brazil where it has been found in all different biomes in the country including areas with high human traffic. [10]
Further information needs to be collected in order to understand the preferable natural environment G. margarita inhabits.
An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualistic relationship. Examples are nitrogen-fixing bacteria, which live in the root nodules of legumes, single-cell algae inside reef-building corals, and bacterial endosymbionts that provide essential nutrients to insects.
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, the plant root system and its surroundings. Mycorrhizae play important roles in plant nutrition, soil biology, and soil chemistry.
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.
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 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.
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.
Fungivory or mycophagy is the process of organisms consuming fungi. Many different organisms have been recorded to gain their energy from consuming fungi, including birds, mammals, insects, plants, amoebas, gastropods, nematodes, bacteria and other fungi. Some of these, which only eat fungi, are called fungivores whereas others eat fungi as only part of their diet, being omnivores.
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.
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.
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.
Fungal-bacterial endosymbiosis encompasses the mutualistic relationship between a fungus and intracellular bacteria species residing within the fungus. Many examples of endosymbiotic relationships between bacteria and plants, algae and insects exist and have been well characterized, however fungal-bacteria endosymbiosis has been less well described.
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.
Candidatus "Glomeribacter gigasporarum" is a gram-negative β-proteobacteria. The bacterium is rod-shaped, and has a obligate endosymbiotic relationship with the arbuscular-mycorrhizal fungi Gigaspora margarita. Sequencing of the16S rRNA gene places Ca. "G. gigasporarum" within the Burkholderia genus. Ca. "G. gigasporarum is unculturable as of yet, but can stay alive in enrichment for up to 4 weeks. The candidate bacteria is considered "the smallest beta-proteobacterium" with a genome size of 1.4 Mb. The chromosome is 750 kb long and a plasmid is 600 to 650 kb. The genome size was determined using gel-electrophoresis.
Dr. Mohamed Hijri is a biologist who studies arbuscular mycorrhizal fungi (AMF). He is a professor of biology and research at the Institut de recherche en biologie végétale at the University of Montreal.
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.
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.
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 Tulasne 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.
The common symbiosis signaling pathway (CSSP) is a signaling cascade in plants that allows them to interact with symbiotic microbes. It corresponds to an ancestral pathway that plants use to interact with arbuscular mycorrhizal fungi (AMF). It is known as "common" because different evolutionary younger symbioses also use this pathway, notably the root nodule symbiosis with nitrogen-fixing rhizobia bacteria. The pathway is activated by both Nod-factor perception, as well as by Myc-factor perception that are released from AMF. The pathway is distinguished from the pathogen recognition pathways, but may have some common receptors involved in both pathogen recognition as well as CSSP. A recent work by Kevin Cope and colleagues showed that ectomycorrhizae also uses CSSP components such as Myc-factor recognition.