Mushrooms: A Kingdom Of Their Own

Mushrooms are the fleshy, spore-bearing fruiting bodies of fungi, typically produced above ground on soil or another food source. The term mushroom is used to identify edible fungi, while toadstool is reserved for inedible or poisonous ones. The standard for the name mushroom is the cultivated white button mushroom, Agaricus bisporus, which has a stem (stipe), a cap (pileus), and gills (lamellae) on the underside of the cap. Mushrooms belong to the Eumycota (true fungi or Eumycetes) group, which includes over 70,000 described species. They are classified into six accepted phylums, including Ascomycota, Basidiomycota, and Zygomycota.

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The kingdom Fungi

Mushrooms belong to the kingdom Fungi, which includes about 144,000 known species of organisms, including yeasts, rusts, smuts, mildews, molds, and mushrooms. Fungi are eukaryotic organisms, meaning their cells contain membrane-bound organelles and clearly defined nuclei. They are distinct from plants and animals, having diverged from these kingdoms around one billion years ago, at the start of the Neoproterozoic Era. Fungi are also structurally different from plants and bacteria, as they have chitin in their cell walls.

Fungi were once considered plant-like organisms, but DNA comparisons have shown that they are more closely related to animals. They are also incapable of photosynthesis and instead acquire their food by absorbing dissolved molecules, secreting digestive enzymes into their environment. They play an essential role in ecosystems as decomposers, breaking down organic materials into simple molecules, and participating in the cycling of nutrients.

Mushrooms, the conspicuous umbrella-shaped fruiting bodies of certain fungi, are typically of the order Agaricales in the phylum Basidiomycota, but they can also belong to other groups. The term "mushroom" is used to identify edible fungi, while "toadstool" is often reserved for inedible or poisonous ones. However, there is no scientific distinction between the two names, and either can be properly applied to any fleshy fungus fruiting structure.

Phylum Basidiomycota

Basidiomycota is a large and diverse phylum of fungi (kingdom Fungi) that includes mushrooms, puffballs, stinkhorns, jelly and shelf fungi, certain yeasts, and the rusts and smuts. Basidiomycota are filamentous fungi composed of hyphae (except for basidiomycota yeast). They reproduce sexually via the formation of specialized club-shaped end cells called basidia that normally bear four external meiospores. These specialized spores are called basidiospores. However, some Basidiomycota reproduce asexually. Basidiomycota that reproduce asexually can typically be recognized as members of this division by gross similarity to others.

Basidiomycota includes agarics, puffballs, stinkhorns, bracket fungi, other polypores, jelly fungi, boletes, chanterelles, earth stars, smuts, bunts, rusts, mirror yeasts, and Cryptococcus, the human pathogenic yeast. Basidiomycota is one of two large divisions that, together with the Ascomycota, constitute the subkingdom Dikarya (often referred to as the "higher fungi") within the kingdom Fungi. Members are known as basidiomycetes.

Basidiomycota were traditionally divided into two classes, now obsolete: Homobasidiomycetes (or holobasidiomycetes), including true mushrooms, and Heterobasidiomycetes, including the jelly, rust, and smut fungi. These former concepts continue to be used as two types of growth habit groupings, the "mushrooms" (e.g. Schizophyllum commune) and the non-mushrooms (e.g. Mycosarcoma maydis). The Agaricomycotina includes what had previously been called the Hymenomycetes, an obsolete morphological class of Basidiomycota that formed hymenial layers on their fruit bodies, the Gasteromycetes, another obsolete class that included species mostly lacking hymenia and mostly forming spores in enclosed fruit bodies, as well as most of the jelly fungi.

Basidiomycota typically have mutually indistinguishable, compatible haploids which are usually mycelia composed of filamentous hyphae. Haploid Basidiomycota mycelia fuse via plasmogamy, and then the compatible nuclei migrate into each other's mycelia and pair up with the resident nuclei. Karyogamy is delayed, so that the compatible nuclei remain in pairs, called dikaryons. The hyphae are then said to be dikaryotic. The dikaryotic mycelium is often more vigorous than individual monokaryotic mycelia and proceeds to take over the substrate in which they are growing. The dikaryons can be long-lived, lasting years, decades, or centuries.

Ascomycota and Basidiomycota

The term "mushroom" is most often used to refer to fungi that have a stem (stipe), a cap (pileus), and gills (lamellae) on the underside of the cap. Most mushrooms are basidiomycetes and gilled, and their spores, called basidiospores, are produced on the gills. The standard for the name "mushroom" is the cultivated white button mushroom, Agaricus bisporus.

Mushrooms belong to the kingdom Fungi, which contains five major phyla that were established according to their mode of sexual reproduction or using molecular data. Most mushrooms belong to the Basidiomycota phylum. Fungi that do not have a known sexual reproductive cycle are presently members of two phyla: Ascomycota and Basidiomycota.

Ascomycota

Ascomycota, or sac fungi, form spores in sacs called asci during sexual reproduction. Asexual reproduction is their most common form of reproduction. Ascomycota includes lichens, and the non-reproductive (vegetative) mycelium of most ascomycetes is usually inconspicuous because it is commonly embedded in the substrate, such as soil, or grows on or inside a living host, and only the ascoma may be seen when fruiting. Many ascomycetes are useful to humans as sources of medicinally important compounds such as antibiotics, as well as for fermenting bread, alcoholic beverages, and cheese.

Basidiomycota

Basidiomycota, or club fungi, produce showy fruiting bodies that contain basidia in the form of clubs. Spores are stored in the basidia. Most mushrooms belong to this phylum.

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Glomeromycota

The fossil record of Glomeromycota is believed to be ancient, extending well back into the Paleozoic. Spores and hyphae of a glomeromycotan type have been reported from rocks as old as the Cambrian and the Upper Ordovician. Palaeoglomus grayi has aseptate (coenocytic) hyphae and spores that resemble living Glomus spores.

The classification of Glomeromycota has been redefined with the adoption of molecular techniques. Initial studies of Glomeromycota were based on the morphology of soil-borne sporocarps (spore clusters) found in or near colonized plant roots. Distinguishing features such as wall morphologies, size, shape, colour, hyphal attachment, and reaction to staining compounds allowed a phylogeny to be constructed. With the advent of molecular techniques, the phylogenetic classification of Glomeromycota has become clearer.

Fungi taxonomy

Historically, fungi were included in the plant kingdom due to similarities in lifestyle and morphology. However, since the 1990s, dramatic changes have occurred in the classification of fungi due to improved understanding and the incorporation of molecular genetics and DNA analysis. Fungi are now recognised as a separate kingdom, the kingdom Fungi, which comprises seven phyla: Chytridiomycota, Blastocladiomycota, Neocallimastigomycota, Microsporidia, Glomeromycota, Ascomycota, and Basidiomycota.

The phylum Chytridiomycota includes the ancestral fungi, while the Microsporidia may also be an ancient sister group. The loss of the chytrid flagellum and the development of branching, aseptate fungal filaments marked the first major steps in the evolution of higher fungi. Septate filaments evolved as Glomeromycota diverged from a combined clade of pre-basidiomycota and pre-ascomycota fungi.

Fungal taxonomy can be challenging due to the simple body plan of fungi and the presence of fungus-like organisms that do not belong to the kingdom Fungi but are often called fungi, such as slime molds and water molds (Oomycota). Species of fungi may be distinguished by their morphological, biochemical, and physiological characteristics, such as spore colour, size and shape, as well as their ability to metabolise certain biochemicals. Molecular identification and sequence-based DNA barcoding are becoming increasingly important approaches in fungal taxonomy, contributing to our understanding of fungal diversity.

The study of fungi is known as mycology, and it involves examining their genetic and biochemical properties, taxonomy, and their use to humans, such as for medicine, food, and religious purposes. The International Commission on the Taxonomy of Fungi (ICTF) and the Nomenclature Committee for Fungi (NCF) play important roles in promoting sound taxonomic practices and considering nomenclatural matters related to fungi.

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