Instead, they are formed by large cells containing many nuclei (multinucleate), an arrangement described as coenocytic hyphae (Figure 4b). Hyphae can form a tangled network called a mycelium and form the thallus (body) of fleshy fungi. Fungi, latin for mushroom, are eukaryotes which are responsible for decomposition and nutrient cycling through the environment. He has published more than 15 research articles and book chapters in international journals and well-renowned publishers. Examples include the blue color in cheeses such as Stilton or Roquefort, which are made by inoculation with Penicillium roqueforti. Somatic cells in yeast form buds. Factors that likely contribute to the under-representation of fungal species among fossils include the nature of fungal fruiting bodies, which are soft, fleshy, and easily degradable tissues and the microscopic dimensions of most fungal structures, which therefore are not readily evident. They display two distinct morphological stages: the vegetative and reproductive. Oenology, the science of wine making, relies not only on the knowledge of grape varietals and soil composition, but also on a solid understanding of the characteristics of the wild yeasts that thrive in different wine-making regions. Fungi store glycogen for their energy needs and use glucose and maltose for . 2018. The most common mode of asexual reproduction is through the formation of asexual spores, which are produced by a single individual thallus (through mitosis) and are genetically identical to the parent thallus (Figure 7). In addition, their mode of nutrition was poorly understood. Fungi are mostly saprobes (saprophyte is an equivalent term): organisms that derive nutrients from decaying organic matter. [42] This process might bear similarity to CO2 fixation via visible light, but instead uses ionizing radiation as a source of energy.[83]. The field of phytopathology, the study of plant diseases, is closely related because many plant pathogens are fungi. Modern penicillins are semisynthetic compounds, obtained initially from fermentation cultures, but then structurally altered for specific desirable properties. [117] Researchers study compression fossils by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details. Other species, such as the Chytridiomycota that reside in the rumen of cattle, are are obligate anaerobes, in that they only use anaerobic respiration because oxygen will disrupt their metabolism or kill them. In common with some plant and animal species, Some species grow as unicellular yeasts that reproduce by, This page was last edited on 17 August 2023, at 17:41. @. The spores of most of the researched species of fungi are transported by wind. We would like to take this opportunity to request your COMMENTS on the topics we covered. Fungi cannot synthesize their own carbohydrates using, CO2, water and sunlight. Mushroom farming and mushroom gathering are large industries in many countries. Instead, they are formed by large cells containing many nuclei, an arrangement described as coenocytic hyphae (Figure 24.1.4). Yet others bud off the vegetative parent cell. 2. Although humans have used yeasts and mushrooms since prehistoric times, until recently, the biology of fungi was poorly understood. Fungi also cause serious infections in plants and animals. [167], Unlike true fungi, the cell walls of oomycetes contain cellulose and lack chitin. Basidiomycota are typically filamentous fungi composed of hyphae. They are described as perforated septa. Introduction to Population Genetics and Speciation, 31. [51] It recognizes seven phyla, two of whichthe Ascomycota and the Basidiomycotaare contained within a branch representing subkingdom Dikarya, the most species rich and familiar group, including all the mushrooms, most food-spoilage molds, most plant pathogenic fungi, and the beer, wine, and bread yeasts. It is possible to purchase yeast strains isolated from specific grape-growing regions. During cell division, the formation of the hook ensures proper distribution of the newly divided nuclei into the apical and basal hyphal compartments. Fungi are more closely related to animals than plants. [16] The Species Survival Commission (SSC) of the International Union for Conservation of Nature (IUCN) in August 2021 asked that the phrase fauna and flora be replaced by fauna, flora, and funga.[17]. [194], Many insects also engage in mutualistic relationships with fungi. [87] Deuteromycota (alternatively known as Deuteromycetes, conidial fungi, or mitosporic fungi) is not an accepted taxonomic clade and is now taken to mean simply fungi that lack a known sexual stage. Fungi are eukaryotic, spore bearing, achlorophyllous, heterotrophic organisms that generally reproduce sexually and asexually and whose filamentous, branched somatic structures are typically surrounded by cell walls containing chitin or cellulose or both with many organic molecules and exhibiting absorptive nutrition. [279][280], Certain fungi, in particular white-rot fungi, can degrade insecticides, herbicides, pentachlorophenol, creosote, coal tars, and heavy fuels and turn them into carbon dioxide, water, and basic elements. Hyphochytrids have both chitin and cellulose. Fungi are a kingdom of multicellular eukaryotic organisms that are heterotrophs (cannot make their own food and rely on other sources) and play key roles in nutrient recycling in an ecosystem. 53. [95] A basidiocarp is formed in which club-like structures known as basidia generate haploid basidiospores after karyogamy and meiosis. [137][138], Some time after the PermianTriassic extinction event (251.4Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in sediments) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available fossil record for this period. Other asexual spores originate in the fragmentation of a hypha to form single cells that are released as spores; some of these have a thick wall surrounding the fragment. The vegetative body of the fungi may be unicellular or composed of microscopic threads called hyphae. Fungi are eukaryotes, and as such, have a complex cellular organization. [38] Mycotoxins may provide fitness benefits in terms of physiological adaptation, competition with other microbes and fungi, and protection from consumption (fungivory). Mycology is a branch of microbiology, and many mycologists start their careers with a degree in microbiology. [102] For example, the structure of the spore-bearing cells in some ascomycete species is such that the buildup of substances affecting cell volume and fluid balance enables the explosive discharge of spores into the air. The Latin adjectival form of "mycology" (mycologic) appeared as early as 1796 in a book on the subject by Christiaan Hendrik Persoon. Alternatively, Rozella can be classified as a basal fungal group. The most common mode of asexual reproduction is through the formation of asexual spores, which are produced by one parent only (through mitosis) and are genetically identical to that parent (Figure \(\PageIndex{7}\)). Fungi thrive in environments that are moist and slightly acidic, and can grow with or without light. Fungi grow best in acidic environment (tolerate acidic pH). [147], There is no unique generally accepted system at the higher taxonomic levels and there are frequent name changes at every level, from species upwards. [32] Hyphae can be either septate or coenocytic. The kingdom Fungi includes an enormous variety of living organisms collectively referred to as Ascomycota, or true Fungi. Many ascomycetes and basidiomycetes go through a dikaryotic stage, in which the nuclei inherited from the two parents do not combine immediately after cell fusion, but remain separate in the hyphal cells (see heterokaryosis). In addition, fungi do not fix nitrogen from the atmosphere. [139] However, the relative proportion of fungal spores relative to spores formed by algal species is difficult to assess,[140] the spike did not appear worldwide,[141][142] and in many places it did not fall on the PermianTriassic boundary. 2023 Microbe Notes. A fungus ( PL: fungi [2] or funguses [3]) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. Fungi are the principal decomposers in ecological systems. fungus, plural fungi, any of about 144,000 known species of organisms of the kingdom Fungi, which includes the yeasts, rusts, smuts, mildews, molds, and mushrooms.There are also many funguslike organisms, including slime molds and oomycetes (water molds), that do not belong to kingdom Fungi but are often called fungi. The great French chemist and microbiologist, Louis Pasteur, made many of his essential discoveries working on the humble brewers yeast, thus discovering the process of fermentation. [169][170], Although often inconspicuous, fungi occur in every environment on Earth and play very important roles in most ecosystems. Structure of Fungi Made with by Sagar Aryal. Fungi digest the food first and then ingest the food, to accomplish this the fungi produce exoenzymes like Hydrolases, Lyases, Oxidoreductase, Transferase, etc. Sexual spores are Oospores, Zygospores, Ascospores, Basidiospores, etc. The genus Blastocystis, now in Stramenopiles, was originally classified as a yeast. [92], Most fungi have both a haploid and a diploid stage in their life cycles. [264] Molds used in cheese production are non-toxic and are thus safe for human consumption; however, mycotoxins (e.g., aflatoxins, roquefortine C, patulin, or others) may accumulate because of growth of other fungi during cheese ripening or storage. Most fungal hyphae are divided into separate cells by endwalls called septa (singular, septum) (Figure \(\PageIndex{4}\)). An ascus (plural asci) is then formed, in which karyogamy (nuclear fusion) occurs. [163], Members of the Basidiomycota, commonly known as the club fungi or basidiomycetes, produce meiospores called basidiospores on club-like stalks called basidia. However, little is known of the true biodiversity of the fungus kingdom, which has been estimated at 2.2million to 3.8million species. 5. also refers to mycology as the study of fungi. Based on fossil evidence, fungi appeared in the pre-Cambrian era, about 450 million years ago. [282][283][284], Several pivotal discoveries in biology were made by researchers using fungi as model organisms, that is, fungi that grow and sexually reproduce rapidly in the laboratory. The wall protects the cell from desiccation and predators. [231] This process is referred to as monokaryotic fruiting. Fungi are heterotrophic organisms. Saccharomyces cerevisiae (bakers yeast) and Candida species (the agents of thrush, a common fungal infection) are examples of unicellular fungi (Figure 2). The alcohol produced from yeast fermentation is used in wine and beer production. and Asexual spores are Sporangiospores, Aplanospores, Zoospores, Conidia, etc. Formation of Organic Molecules in an Earthly Reducing Atmosphere, 71. Smut and Dutch elm disease affect plants, whereas athletes foot and candidiasis (thrush) are medically important fungal infections in humans. General characters of fungi Fungi are the eukaryotic, achlorophyllous, and unicellular or multicellular organisms, which may reproduce by asexual and sexual spores. Under some current systematic phylogenies, they continue to be a polyphyletic group of organisms that share characteristics, rather than sharing a single common ancestor. [209] This same principle is applied to fungi-feeding parasites, including Asterotremella albida, which feeds on the fruit bodies of other fungi both while they are living and after they are dead. During karyogamy (nuclear marriage), the haploid nuclei fuse to form a diploid zygote nucleus. Fungi are a kingdom of usually multicellular eukaryotic organisms that are heterotrophs (cannot make their own food) and have important roles in nutrient cycling in an ecosystem. Other asexual spores originate in the fragmentation of a hypha to form single cells that are released as spores; some of these have a thick wall surrounding the fragment. To feed, both types of fungi secrete digestive, Some parasitic fungi also produce special absorptive organs called. Fungi. For example, the giant puffball mushroom bursts open and releases trillions of spores in a massive cloud of what looks like finely particulate dust. Fungi often interact with other organisms, forming beneficial or mutualistic associations. [10], The word mycology is derived from the Greek mykes ( 'mushroom') and logos ( 'discourse'). There are many types of asexual spores. Want to create or adapt books like this? Mycologists collaborate with zoologists and plant pathologists to identify and control difficult fungal infections, such as the devastating chestnut blight, the mysterious decline in frog populations in many areas of the world, or the deadly epidemic called white nose syndrome, which is decimating bats in the Eastern United States. [72] Adaptations such as these are complemented by hydrolytic enzymes secreted into the environment to digest large organic moleculessuch as polysaccharides, proteins, and lipidsinto smaller molecules that may then be absorbed as nutrients. During budding (a type of cytokinesis), a bulge forms on the side of the cell, the nucleus divides mitotically, and the bud ultimately detaches itself from the mother cell (Figure \(\PageIndex{6}\)). These colonies can exhibit growth shapes and colors (due to spores or pigmentation) that can be used as diagnostic features in the identification of species or groups. The shiitake mushroom is a source of lentinan, a clinical drug approved for use in cancer treatments in several countries, including Japan. The fungal part of the relationship is composed mostly of various species of ascomycetes and a few basidiomycetes. Unicellular fungi are generally referred to as yeasts. [227] Furthermore, U. maydis has a well-established recombinational DNA repair system which acts during mitosis and meiosis. Nonetheless, studies of the oomycetes and myxomycetes are still often included in mycology textbooks and primary research literature.[168]. [51][146] The taxonomy of fungi is in a state of constant flux, especially due to research based on DNA comparisons. [216], Organisms that parasitize fungi are known as mycoparasitic organisms. For example, Dutch elm disease, which is caused by the fungus Ophiostoma ulmi, is a particularly devastating type of fungal infestation that destroys many native species of elm (Ulmus sp.) They are classified as heterotrophs among the living organisms. Thank you the information is very helpful, Very helpful can send some summaries about the kingdom animalia, Send me notes for, the details structure of the cell membrane. Most fungi are multicellular organisms. [214] Other fungi can attack eyes, nails, hair, and especially skin, the so-called dermatophytic and keratinophilic fungi, and cause local infections such as ringworm and athlete's foot. [121], The Neocallimastigomycota were earlier placed in the phylum Chytridiomycota. [57], Most fungi grow as hyphae, which are cylindrical, thread-like structures 210m in diameter and up to several centimeters in length. [70] The pressure generated by the appressorium, directed against the plant epidermis, can exceed 8 megapascals (1,200psi). Because fungi feed on decaying and dead matter, they are saprobes. The carbon, nitrogen, and other elements are thus released into the environment. As eukaryotes, fungi possess a biosynthetic pathway for producing terpenes that uses mevalonic acid and pyrophosphate as chemical building blocks. Extensions of the Laws of Inheritance, 26. In humans, fungal infections are generally considered challenging to treat. For example, some species of fungi can be used to break down diesel oil and polycyclic aromatic hydrocarbons (PAHs). Most fungi grow at an acidic pH of about 5.0, although some species grow at lower and higher pH levels. As decomposers, they play an essential role in nutrient cycling, especially as saprotrophs and symbionts, degrading organic matter to inorganic molecules, which can then re-enter anabolic metabolic pathways in plants or other organisms. They obtain their nutrients from dead or decomposing organic matter: mainly plant material. Historically, fly agaric was used by different peoples in Europe and Asia and its present usage for religious or shamanic purposes is reported from some ethnic groups such as the Koryak people of northeastern Siberia. They have long been used as a direct source of human food, in the form of mushrooms and truffles; as a leavening agent for bread; and in the fermentation of various food products, such as wine, beer, and soy sauce. They are often used in gourmet dishes. This phylum includes morels, a few mushrooms and truffles, unicellular yeasts (e.g., of the genera Saccharomyces, Kluyveromyces, Pichia, and Candida), and many filamentous fungi living as saprotrophs, parasites, and mutualistic symbionts (e.g. They are eukaryotes that have cell walls, are heterotrophs that feed by absorbing their food, and use spores to reproduce. Unlike bacteria, fungi do not respond to traditional antibiotic therapy because they are eukaryotes. All Rights Reserved. In fact, up until the mid-20th century, many scientists classified fungi as plants! Origins of Organic Molecules in a Non-Reducing Atmosphere, 72. Although inaccurate, the common misconception that fungi are plants persists among the general public due to their historical classification, as well as several similarities. Progress in the field of fungal biology was the result of mycology: the scientific study of fungi. Fungi, like plants, arose mostly sessile and seemingly rooted in place. They are non-vascular organisms. A few types of fungi have accessory genomic structures comparable to bacterial plasmids (loops of DNA); however, the horizontal transfer of genetic information that occurs between one bacterium and another rarely occurs in fungi. Nutrition in fungi they are saprophytes (gets energy from dead and decaying matters), or parasites (lives in a host, attack and kill) or symbionts (mutually beneficial). [93], In ascomycetes, dikaryotic hyphae of the hymenium (the spore-bearing tissue layer) form a characteristic hook (crozier) at the hyphal septum. They range in size from unicellular yeasts to large multicellular fungi. By the end of this section, you will be able to do the following: Although humans have used yeasts and mushrooms since prehistoric times, until recently, the biology of fungi was poorly understood. Is the Brain Another Object of Sexual Desire? Although individual hyphae must be observed under a microscope, the mycelium of a fungus can be very large, with some species truly being the fungus humongous. The giant Armillaria solidipes (honey mushroom) is considered the largest organism on Earth, spreading across more than 2,000 acres of underground soil in eastern Oregon; it is estimated to be at least 2,400 years old. Describe the basic characteristics of fungi. As with animal cells, the polysaccharide of storage is glycogen, a branched polysaccaride, rather than amylopectin, a less densely branched polysaccharide, and amylose, a linear polysaccharide, as found in plants. The poisonous Amanita muscaria (fly agaric) is recognizable by its bright red cap with white patches (Figure \(\PageIndex{1}\)). Fungi are eukaryotes, and as such, have a complex cellular organization. Other fungal models have emerged that address specific biological questions relevant to medicine, plant pathology, and industrial uses; examples include Candida albicans, a dimorphic, opportunistic human pathogen,[286] Magnaporthe grisea, a plant pathogen,[287] and Pichia pastoris, a yeast widely used for eukaryotic protein production. Progress in the field of fungal biology was the result of mycology: the scientific study of fungi. 55. This page titled 24.1: Characteristics of Fungi is shared under a CC BY 4.0 license and was authored, remixed, and/or curated by OpenStax. Morphological characteristics of fungi:mycelium and hyphae . For example, certain species eliminate or suppress the growth of harmful plant pathogens, such as insects, mites, weeds, nematodes, and other fungi that cause diseases of important crop plants. Fungi, once considered plant-like organisms, are more closely related to animals than plants. Spores allow fungi to expand their distribution and colonize new environments. Soil rich in organic matter is an ideal habitat for many species, and only a small number of fungi are found in drier areas or in habitats with little or no organic matter. They are described as perforated septa. When the zygospore germinates, it undergoes meiosis, generating new haploid hyphae, which may then form asexual sporangiospores. Mostly they are terrestrial, parasitic or coprophilous. Conidiospores are unicellular or multicellular spores that are released directly from the tip or side of the hypha. This electron micrograph shows (c) the spore-bearing structures of Aspergillus, a type of toxic fungi found mostly in soil and plants. Many fungi display bright colors arising from other cellular pigments, ranging from red to green to black. To overcome this limitation, some fungi, such as Armillaria, form rhizomorphs,[37] which resemble and perform functions similar to the roots of plants. Fungi are the source of many commercial enzymes and antibiotics. They are therefore chemoheterotrophic organisms. When both mating types are present in the same mycelium, it is called homothallic, or self-fertile. Karyogamy in the asci is followed immediately by meiosis and the production of ascospores. Another site for additional information and photographs: A short video on fungi - this is a short Youtube - skip the ad when it comes on and the information on fungi starts after 1 minute - advance to skip the intro photos. Multicellular fungi produce threadlike hyphae (singular hypha). A specialized anatomical structure, called a clamp connection, is formed at each hyphal septum. [118], The earliest fossils possessing features typical of fungi date to the Paleoproterozoic era, some 2,400million years ago (Ma); these multicellular benthic organisms had filamentous structures capable of anastomosis. [103] The forcible discharge of single spores termed ballistospores involves formation of a small drop of water (Buller's drop), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000g;[104] the net result is that the spore is ejected 0.010.02cm, sufficient distance for it to fall through the gills or pores into the air below. [89] The symbiotic association between the Glomeromycota and plants is ancient, with evidence dating to 400million years ago. Furthermore, persons with immuno-deficiencies are particularly susceptible to disease by genera such as Aspergillus, Candida, Cryptoccocus,[179][211][212] Histoplasma,[213] and Pneumocystis. Introductory Biology: Evolutionary and Ecological Perspectives, Review the characteristics of fungi by visiting this interactive site from Wisconsin-online, https://openstax.org/books/biology-2e/pages/1-introduction, Explain sexual and asexual reproduction in fungi. Fungi include symbionts of plants, animals, or other fungi and also parasites. Some fungi are macroscopic and can be seen by naked eyes. [154] Members of the group that have been isolated lack the chitinous cell wall that is characteristic of fungi. In fungi, sexual reproduction often occurs in response to adverse environmental conditions. [36], Fungal mycelia can become visible to the naked eye, for example, on various surfaces and substrates, such as damp walls and spoiled food, where they are commonly called molds. [217] Fungi can also act as mycoparasites or antagonists of other fungi, such as Hypomyces chrysospermus, which grows on bolete mushrooms. The great French chemist and microbiologist, Louis Pasteur, made many of his essential discoveries working on the humble brewers yeast, thus discovering the process of fermentation. Wrapping Up: Understanding the Silent Crickets, 61. Mycologists collaborate with zoologists and plant pathologists to identify and control difficult fungal infections, such as the devastating chestnut blight, the mysterious decline in frog populations in many areas of the world, or the deadly epidemic called white nose syndrome, which is decimating bats in the Eastern United States. Fungi reproduce both sexually and asexually, and they also have symbiotic associations with plants and bacteria. Sexual reproduction involves plasmogamy (the fusion of the cytoplasm), followed by karyogamy (the fusion of nuclei). Reproduction in fungi is both by sexual and asexual means. Serious pathogens of many cultivated plants causing extensive damage and losses to agriculture and forestry include the rice blast fungus Magnaporthe oryzae,[205] tree pathogens such as Ophiostoma ulmi and Ophiostoma novo-ulmi causing Dutch elm disease,[206] Cryphonectria parasitica responsible for chestnut blight,[207] and Phymatotrichopsis omnivora causing Texas Root Rot, and plant pathogens in the genera Fusarium, Ustilago, Alternaria, and Cochliobolus. Jun 8, 2022 24.1A: Characteristics of Fungi 24.1C: Fungi Reproduction Boundless Boundless Fungi are unicellular or multicellular thick-cell-walled heterotroph decomposers that eat decaying matter and make tangles of filaments. Some fungal pigments are toxic to humans. Two amber-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species Archaeomarasmius leggetti) appeared during the late Cretaceous, 90Ma. However, there are also many fungi species that dont produce mushrooms at all. 56. Fungi grow best in acidic environment (tolerate acidic pH). Genera with mushrooms containing deadly toxins include Conocybe, Galerina, Lepiota and the most infamous, Amanita. They are heterotrophs and contain neither photosynthetic pigments such as chlorophyll, nor organelles such as chloroplasts. Determining Evolutionary Relationships, 34. Most fungi are obligate aerobes, requiring oxygen to survive. Yeasts are intermediate, being facultative anaerobes. The fungus then penetrates the tissue of the worm by extending specialized hyphae called haustoria. [5] The number of new fungi species discovered yearly has increased from 1,000 to 1,500 per year about 10 years ago, to about 2000 with a peak of more than 2,500 species in 2016. Fungal spores are smaller and lighter than plant seeds. Fungi reproduce via spores. The bacteria were also included in fungi in some classifications, as the group Schizomycetes. Growth rate of fungi is slower than that of bacteria. Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their cryptic lifestyles in soil or on dead matter. [136] Rare in the fossil record are the Homobasidiomycetes (a taxon roughly equivalent to the mushroom-producing species of the Agaricomycetes). Like plant cells, fungal cells have a thick cell wall. The cell wall is made up of chitin or -glucans. Language links are at the top of the page across from the title. Examples of statins found in fungi include mevastatin from Penicillium citrinum and lovastatin from Aspergillus terreus and the oyster mushroom. Asci are embedded in an ascocarp, or fruiting body. However, the dikaryotic phase is more extensive in the basidiomycetes, often also present in the vegetatively growing mycelium. Sexual reproduction in basidiomycetes is similar to that of the ascomycetes. The application of molecular tools, such as DNA sequencing and phylogenetic analysis, to study diversity has greatly enhanced the resolution and added robustness to estimates of genetic diversity within various taxonomic groups. Pigments in fungi are associated with the cell wall and play a protective role against ultraviolet radiation. document.getElementById( "ak_js_1" ).setAttribute( "value", ( new Date() ).getTime() ); This site uses Akismet to reduce spam. They display two distinct morphological stages: the vegetative and reproductive. External enzymes digest nutrients that are absorbed by the body of the fungus, which is called a thallus. 11. Many species produce metabolites that are major sources of pharmacologically active drugs. [255], Baker's yeast or Saccharomyces cerevisiae, a unicellular fungus, is used to make bread and other wheat-based products, such as pizza dough and dumplings. Dimorphic fungi can change from the unicellular to multicellular state depending on environmental conditions. [46], As of 2020,[update] around 148,000 species of fungi have been described by taxonomists,[6] but the global biodiversity of the fungus kingdom is not fully understood. These organisms are classified as a kingdom,[4] separately from the other eukaryotic kingdoms, which, by one traditional classification, includes Plantae, Animalia, Protozoa, and Chromista. For example, some species of fungi can be used to break down diesel oil and polycyclic aromatic hydrocarbons (PAHs). As with the structurally similar hook in the ascomycetes, the clamp connection in the basidiomycetes is required for controlled transfer of nuclei during cell division, to maintain the dikaryotic stage with two genetically different nuclei in each hyphal compartment.
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