Mycelium

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Mycelium
Various examples of mycelium in different sizes, environments and species.

Mycelium (pl.: mycelia) [lower-alpha 1] is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. [1] Its normal form is that of branched, slender, entangled, anastomosing, hyaline threads. [2] Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates into a monokaryotic mycelium, [1] which cannot reproduce sexually; when two compatible monokaryotic mycelia join and form a dikaryotic mycelium, that mycelium may form fruiting bodies such as mushrooms. [3] A mycelium may be minute, forming a colony that is too small to see, or may grow to span thousands of acres as in Armillaria .

Contents

Through the mycelium, a fungus absorbs nutrients from its environment. It does this in a two-stage process. First, the hyphae secrete enzymes onto or into the food source, which break down biological polymers into smaller units such as monomers. These monomers are then absorbed into the mycelium by facilitated diffusion and active transport.

Mycelia are vital in terrestrial and aquatic ecosystems for their role in the decomposition of plant material. They contribute to the organic fraction of soil, and their growth releases carbon dioxide back into the atmosphere (see carbon cycle). Ectomycorrhizal extramatrical mycelium, as well as the mycelium of arbuscular mycorrhizal fungi, increase the efficiency of water and nutrient absorption of most plants and confers resistance to some plant pathogens. Mycelium is an important food source for many soil invertebrates. They are vital to agriculture and are important to almost all species of plants, many species co-evolving with the fungi. Mycelium is a primary factor in some plants’ health, nutrient intake and growth, with mycelium being a major factor to plant fitness.

Networks of mycelia can transport water [4] and spikes of electrical potential. [5]

Sclerotia are compact or hard masses of mycelium.

Uses

Agriculture

One of the primary roles of fungi in an ecosystem is to decompose organic compounds. Petroleum products and some pesticides (typical soil contaminants) are organic molecules (i.e., they are built on a carbon structure), and thereby show a potential carbon source for fungi. Hence, fungi have the potential to eradicate such pollutants from their environment unless the chemicals prove toxic to the fungus. This biological degradation is a process known as bioremediation.

Mycelial mats have been suggested as having potential as biological filters, removing chemicals and microorganisms from soil and water. The use of fungal mycelium to accomplish this has been termed mycofiltration.

Knowledge of the relationship between mycorrhizal fungi and plants suggests new ways to improve crop yields. [6]

When spread on logging roads, mycelium can act as a binder, holding disturbed new soil in place preventing washouts until woody plants can establish roots.

Fungi are essential for converting biomass into compost, as they decompose feedstock components such as lignin, which many other composting microorganisms cannot. [7] Turning a backyard compost pile will commonly expose visible networks of mycelia that have formed on the decaying organic material within. Compost is an essential soil amendment and fertilizer for organic farming and gardening. Composting can divert a substantial fraction of municipal solid waste from landfills. [8]

Commercial

Alternatives to polystyrene and plastic packaging can be produced by growing mycelium in agricultural waste. [9]

Mycelium has also been used as a material in furniture, and artificial leather. [10]

One of the main commercial uses of mycelium is its use to create artificial leather. Animal leather contributes to a significant environmental footprint, as livestock farming is associated with deforestation, greenhouse gas emissions, and grazing. In addition, the production of synthetic leathers from polyvinyl chloride and polyurethane require the use of hazardous chemicals and fossil fuels, and they are not biodegradable (like plastic). Fungal-based artificial leather is cheaper to produce, has less of an environmental footprint, and is biodegradable. It costs between 18 and 28 cents to produce a square meter of raw mycelium, while it costs between $5.81 and $6.24 to produce a square meter of raw animal hide. Fungal growth is carbon neutral and pure mycelium is 94% biodegradable. However, the use of polymeric materials such as polyester or polylactic acid to improve artificial leather’s properties can negatively affect the biodegradability of the material. [11]

To create leather, fungal mycelium is grown either using liquid-state or solid-state fermentation. In liquid-state fermentation, companies typically use laboratory media or agricultural byproducts to grow fungal biomass. The fungal biomass is then separated into fibers and processed using fiber suspension, filtration, pressing, and drying. These techniques are also commonly utilized in traditional papermaking processes. In solid-state fermentation, mycelium is grown on forestry bioproducts, like sawdust, in an environment with high carbon dioxide concentrations and controlled humidity and temperature. The mycelium mat formed on top of the particle bed is dehydrated, chemically treated, and then compressed to a desired thickness and engraved with a pattern. [11]

Construction material

Mycelium is a strong candidate for sustainable construction primarily due to its lightweight biodegradable structure and its capacity to be grown from waste sources. In addition to this, mycelium has a relatively high strength-to-weight ratio and a much lower embodied energy compared to traditional building materials. Because mycelium takes the form of any mold it's grown in, it can also be advantageous for customization purposes, especially if it's employed as an architectural or aesthetic feature. Current research has also indicated that mycelium does not release toxic resins in the event of a fire because it has a charring effect similar to mass timber. Mycelium plays an interesting role in acoustic insulation, boasting of an absorbance of 70–75% for frequencies of 1500 Hz or less. [12]

Strengths and weaknesses

Mycelium bio-composites have shown strong potential for structural applications, with much higher strength-to-weight ratios than that of conventional materials due primarily to its low density. Compared to conventional building materials, mycelium also has a number of desirable properties that make it an attractive alternative. For example, it has low thermal conductivity and can provide high acoustic insulation. It is biodegradable, has much lower embodied energy, and can serve as a carbon sink, which makes mycelium bio-composites a possible solution to the emissions, energy, and waste associated with building construction.

While mycelium proposes interesting implications as a structural material, there are several significant disadvantages that make it difficult to be practically implemented in large-scale projects. For one, mycelium does not have particularly high compressive strength on its own, ranging from 0.1-0.2 MPa. [13] This is in stark comparison to traditional concrete, which typically has a compressive strength of 17-28 MPa. Even more, because mycelium is considered a living material, it holds specific requirements that make it susceptible to environmental conditions. For instance, it requires a constant source of air in order to stay alive, needs a relatively humid habitat to grow, and cannot be exposed to large amounts of water for fear of contamination and decay.

Mechanical properties

3 separate fungi species (Colorius versicolor, Trametes ochracea, and Ganoderma sessile) were mixed independently with 2 substrates (apple and vine) and tested under separate incubation conditions in order to quantify certain mechanical properties of mycelium. In order to do this, samples were grown in molds, incubated, and dried over the course of 12 days. Samples were tested for water absorption using ASTM C272 guidelines and compared against an EPS material. Tiles of uniform size were cut from the fabricated mold and put under an Instron 3345 machine going at 1 mm/min, up until 20% deformation. [14]

Throughout a 4 stage process, the impact of various substrate and fungal mixes was investigated along with properties of mycelium such as density, water absorption, and compressive strength. Samples were separated into two separate incubation methods and inspected for differences in color, texture, and growth. For the same fungi within each incubation method, minimal differences were recorded. However, across disparate substrate mixtures within the same fungi, colorization and external growth varied between the test samples. While loss of organic matter was calculated, no uniform correlation was found between substrate used and chemical properties of the material. For each of the substrate-fungi mixtures, average densities ranged from 174.1 kg/m3 to 244.9 kg/m3, with the Ganoderma sessile fungi and apple substrate combination being the most dense. Compression tests revealed the Ganoderma sessile fungi and vine substrate to have the highest strength of the samples tested, but no numerical value was provided. [14] For reference, surrounding literature has provided a ballpark estimate of 1-72 kPa. Beyond this, mycelium has a thermal conductivity of 0.05–0.07W/m·K which is less than that of typical concrete. [15]

Construction

The construction of mycelium structures is primarily categorized into three approaches. These include growing blocks in molds, growing in place monolithic structures, and bio-welded units. The first approach cultivates mycelium and its substrate in forms, after which it is dried in ovens and then transported and assembled on site. The second approach uses existing formwork and adapts cast-in-place concrete techniques to grow monolithic mycelium structures in place. The third approach is a hybrid of the previous two referred to as myco-welding, where individual pre-grown units are grown together into a larger monolithic structure. [13]

Studies using grow-in-place methods and myco-welding have explored how to cultivate mycelium and re-use formwork in construction and investigated post-tensioning and friction connections. Research in fabrication has revealed some common challenges faced in construction of mycelium structures, mostly related to the growth of the fungi. It can be difficult to cultivate living material into formwork and it is susceptible to contamination if not properly sterilized. The fungi needs to be kept refrigerated to prevent hardening and properly manage growth and substrate consumption. Additionally, the thickness of fungal growth is limited by the presence of oxygen; if there is no oxygen, the center of the growth can die or be contaminated. [13]

Environmental impact

Researchers have performed life-cycle assessments to evaluate the environmental impact of mycelium bio-composites. Life cycle analysis showed the viability of mycelium as a carbon sink material and as a sustainable alternative to conventional building materials. [12] Use of mycelium as a natural adhesive material may provide environmental benefits, as the fungal-based composites that mycelium is used to create are low cost, low emission, and sustainable. These composites also have a wide range of applications and uses, many of which are in industries responsible for significant environmental pollution, like construction and packaging. [16]

Modern construction and packaging materials are industrially fabricated, non-recyclable, and pollutive: wood products lead to severe deforestation and weather fluctuation; cement is nonbiodegradable and causes high emissions both in production and demolition. Mycelium appears to be cheaper and more sustainable than its counterparts. [16]

Mycelium’s adhesive properties are largely responsible for its diverse array of applications, as it allows them to bind certain substances together. These properties are products of their biological processes, as they secrete corrosive enzymes that allow them to degrade and colonize organic substrates. During degradation, mycelium develops a dense network of thin strands that fuse together within the organic substrate, creating solid material that can hold multiple substrates together. This self-assembly property of mycelium is quite unique, and allows mycelium to grow on a wide range of organic material, including organic waste. [16]

Potential ecological role

Plants appear to communicate within an ecosystem using mycelium, the fungal network produced by mycorrhiza fungi. [17] Mycelial networks constitute 20-30% of soil biomass, though traditional biomass measures fail to detect them. Some 83% of plants appear to exhibit mutualistic association with mycelium as an extension of their root systems, with varying levels of reliance. [18] By some estimates, mycelial networks receive up to 10% or more of the photosynthesis output of their host plants.[ citation needed ]

This mutualism is initiated by hyphal connections in which mycelial strands infect and attach themselves to plant hyphae, penetrating the cell wall but not entering through the membrane into the plant cytoplasm. Mycelium interacts with the cell at the periarbuscular membrane, which behaves as a sort of exchange medium for nutrients and can produce electrical gradients allowing for electrophysiological signals to be sent and received. [17] In modeling studies, different fungi supply different levels of nutrients and growth-promoting materials, with plants tending to root towards (and thus being infected by) fungi supplying most mineral phosphorus and nitrogen (both essential for plant growth). [19]

Mycorrhizal mycelial associations may intensify competition between individuals of the same species, while alleviating competition between species, via the promotion of inferior competitors, thus promoting plant diversity within its network. [17] In doing so, mycorrhizal fungi promote community ecology, with an added complexity of niche differentiation of different networks and types of mycorrhizal fungi that root at different depths, disperse different organic compounds and nutrients, and have unique interactions with specific species of plants. [17]

Mycelial biology and memory

Several studies have documented the memory capacity of mycelial networks and their adaptability to specific environmental conditions. Mycelia have been specialized for different functions in various climates and develop symbiotic or pathogenic relationships with other organisms, such as the human pathogen Candida auris , which has developed a unique approach of evading detection by human neutrophils through adaptive selection–a process of fungal learning and memory. [20] Additionally, these functions can change based on the scale of the mycelia and nature of the symbiotic relationship; commensal and mutual relationships between fungi and plants form through a separate process known as mycorrhizal association, which are called mycorrhiza. Additionally, hyphal organization into mycelial networks can be deterministic for a variety of functions including biomass retention, water recycling, expansion of future hyphae on a resource efficient approach towards desired nutrient gradients, and the subsequent distribution of these resources across the hyphal network. [21] On a macroscopic scale, many mycelia operate with a sort of hierarchy having a “trunk” or main mycelium, with smaller “branches” branching off.  Some saprotrophic basidiomycetes are able to remember past decisions about directional nutrition gradients and will build future mycelium in that direction. [22]

Mycelial memory and intelligence

Current research on collective mycelial intelligence is limited, and while many studies have observed memory and the exchange of electric charge across mycelial networks, this is insufficient evidence to make conclusions about how sensory data is processed in these networks. However, some examples of increased thermal resistance in filamentous fungi suggest a power-law relationship for memory and exposure to a stimulus. [23] Mycelia have also demonstrated the ability to edit their genetic structures within a lifetime due to antibiotic or other extracellular stressors, which can cause rapid acquisition of resistance genes, like those in C. auris . [20] Additionally, plasmodial slime molds demonstrate a similar method of information sharing, as both mycelia and slime molds make use of cAMP molecules for aggregation and signaling. [21]

Sclerotium

Sclerotium is a compact mass of hardened mycelium. For many years, sclerotia were mistaken for individual organisms and described as separate species. However, in the mid 19th century, it was proven that sclerotia was simply a stage in the life cycle of many fungi. Sclerotia are composed of thick, dense shells with dark cells. They are rich in hyphae emergency supplies, such as oil, and they contain small amounts of water. They can survive in dry environments for many years without losing the ability to grow. The size of sclerotia can range from less than a millimeter to tens of centimeters in diameter. [24]

See also

Related Research Articles

<span class="mw-page-title-main">Ascomycota</span> Division or phylum of fungi

Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The defining feature of this fungal group is the "ascus", a microscopic sexual structure in which nonmotile spores, called ascospores, are formed. However, some species of Ascomycota are asexual and thus do not form asci or ascospores. Familiar examples of sac fungi include morels, truffles, brewers' and bakers' yeast, dead man's fingers, and cup fungi. The fungal symbionts in the majority of lichens such as Cladonia belong to the Ascomycota.

<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">Decomposer</span> Organism that breaks down dead or decaying organisms

Decomposers are organisms that break down dead or decaying organisms; they carry out decomposition, a process possible by only certain kingdoms, such as fungi. Like herbivores and predators, decomposers are heterotrophic, meaning that they use organic substrates to get their energy, carbon and nutrients for growth and development. While the terms decomposer and detritivore are often interchangeably used, detritivores ingest and digest dead matter internally, while decomposers directly absorb nutrients through external chemical and biological processes. Thus, invertebrates such as earthworms, woodlice, and sea cucumbers are technically detritivores, not decomposers, since they are unable to absorb nutrients without ingesting them.

<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.

<span class="mw-page-title-main">Sclerotium</span> Mycelial mass

A sclerotium, is a compact mass of hardened fungal mycelium containing food reserves. One role of sclerotia is to survive environmental extremes. In some higher fungi such as ergot, sclerotia become detached and remain dormant until favorable growth conditions return. Sclerotia initially were mistaken for individual organisms and described as separate species until Louis René Tulasne proved in 1853 that sclerotia are only a stage in the life cycle of some fungi. Further investigation showed that this stage appears in many fungi belonging to many diverse groups. Sclerotia are important in the understanding of the life cycle and reproduction of fungi, as a food source, as medicine, and in agricultural blight management.

<span class="mw-page-title-main">Soil biology</span> Study of living things in soil

Soil biology is the study of microbial and faunal activity and ecology in soil. Soil life, soil biota, soil fauna, or edaphon is a collective term that encompasses all organisms that spend a significant portion of their life cycle within a soil profile, or at the soil-litter interface. These organisms include earthworms, nematodes, protozoa, fungi, bacteria, different arthropods, as well as some reptiles, and species of burrowing mammals like gophers, moles and prairie dogs. Soil biology plays a vital role in determining many soil characteristics. The decomposition of organic matter by soil organisms has an immense influence on soil fertility, plant growth, soil structure, and carbon storage. As a relatively new science, much remains unknown about soil biology and its effect on soil ecosystems.

<span class="mw-page-title-main">Saprotrophic nutrition</span> Type of heterotrophic nutrition based on decayed organic matter

Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed organic matter. It occurs in saprotrophs, and is most often associated with fungi and soil bacteria. Saprotrophic microscopic fungi are sometimes called saprobes. Saprotrophic plants or bacterial flora are called saprophytes, although it is now believed that all plants previously thought to be saprotrophic are in fact parasites of microscopic fungi or other plants. In fungi, the process is most often facilitated through the active transport of such materials through endocytosis within the internal mycelium and its constituent hyphae.

<i>Rhizoctonia solani</i> Species of fungus

Rhizoctonia solani is a species of fungus in the order Cantharellales. Basidiocarps are thin, effused, and web-like, but the fungus is more typically encountered in its anamorphic state, as hyphae and sclerotia. The name Rhizoctonia solani is currently applied to a complex of related species that await further research. In its wide sense, Rhizoctonia solani is a facultative plant pathogen with a wide host range and worldwide distribution. It causes various plant diseases such as root rot, damping off, and wire stem. It can also form mycorrhizal associations with orchids.

<span class="mw-page-title-main">Mycelial cord</span> Structure produced by fungi

Mycelial cords are linear aggregations of parallel-oriented hyphae. The mature cords are composed of wide, empty vessel hyphae surrounded by narrower sheathing hyphae. Cords may look similar to plant roots, and also frequently have similar functions; hence they are also called rhizomorphs. As well as growing underground or on the surface of trees and other plants, some fungi make mycelial cords which hang in the air from vegetation.

<span class="mw-page-title-main">Mycoforestry</span> Permaculture forest management system using fungi

Mycoforestry is an ecological forest management system implemented to enhance forest ecosystems and plant communities through the introduction of mycorrhizal and saprotrophic fungi. Mycoforestry is considered a type of permaculture and can be implemented as a beneficial component of an agroforestry system. Mycoforestry can enhance the yields of tree crops and produce edible mushrooms, an economically valuable product. By integrating plant-fungal associations into a forestry management system, native forests can be preserved, wood waste can be recycled back into the ecosystem, carbon sequestration can be increased, planted restoration sites are enhanced, and the sustainability of forest ecosystems are improved. Mycoforestry is an alternative to the practice of clearcutting, which removes dead wood from forests, thereby diminishing nutrient availability and reducing soil depth.

<span class="mw-page-title-main">Nidulariaceae</span> Family of fungi

The Nidulariaceae are a family of fungi in the order Agaricales. Commonly known as the bird's nest fungi, their fruiting bodies resemble tiny egg-filled birds' nests. As they are saprobic, feeding on decomposing organic matter, they are often seen growing on decaying wood and in soils enriched with wood chips or bark mulch; they have a widespread distribution in most ecological regions. The five genera within the family, namely, Crucibulum, Cyathus, Mycocalia, Nidula, and Nidularia, are distinguished from each other by differences in morphology and peridiole structure; more recently, phylogenetic analysis and comparison of DNA sequences is guiding new decisions in the taxonomic organization of this family.

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

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

Mycorrhizal associations have profoundly impacted the evolution of plant life on Earth ever since the initial adaptation of plant life to land. In evolutionary biology, mycorrhizal symbiosis has prompted inquiries into the possibility that symbiosis, not competition, is the main driver of evolution.

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

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.

<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.

<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.

<span class="mw-page-title-main">Mycelium-based materials</span>

Mycelium, a root-like structure that comprises the main vegetative growth of fungi, has been identified as an ecologically friendly substitute to a litany of materials throughout different industries, including but not limited to packaging, fashion and building materials. Such substitutes present a biodegradable alternative to conventional materials.

<span class="mw-page-title-main">Fungi in art</span> Direct and indirect influence of fungi in the arts

Fungi are a common theme and working material in art. Fungi appear in nearly all art forms, including literature, paintings, and graphic arts; and more recently, contemporary art, music, photography, comic books, sculptures, video games, dance, cuisine, architecture, fashion, and design. There are some exhibitions dedicated to fungi, as well as an entire museum.

References

Footnotes

  1. This is the preferred plural form, though mycelium, like fungus, can be considered either singular or plural.

Citations

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