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''Mucor racemosus'' is a rapidly growing, weedy mould belonging to the division
Mucoromycota Mucoromycota is a division within the kingdom fungi. They include a diverse group of various molds, including the common bread molds ''Mucor'' and ''Rhizopus''. It is a sister phylum to Dikarya. It consists of mainly mycorrhizal fungi, root end ...
. It is one of the earliest fungi to be grown in pure culture and was first isolated in 1886. It has a worldwide distribution and colonizes many habitats such as vegetational products, soil and houses. The fungus is mostly known for its ability to exhibit both filamentous and yeast-like morphologies, often referred to as dimorphism. Stark differences are seen in both forms and conditions of the environment heavily affect the phases of the ''M. racemosus''. Like many fungi, it also reproduces both sexually and asexually. The dimorphic capacity of this species has been proposed as an important factor in its pathogenicity and has enhanced the industrial importance. This species is considered an opportunistic pathogen, generally limited to immunocompromised individuals. It also been associated with
allergy Allergies, also known as allergic diseases, refer a number of conditions caused by the hypersensitivity of the immune system to typically harmless substances in the environment. These diseases include hay fever, food allergies, atopic derma ...
and
inflammation Inflammation (from la, wikt:en:inflammatio#Latin, inflammatio) is part of the complex biological response of body tissues to harmful stimuli, such as pathogens, damaged cells, or Irritation, irritants, and is a protective response involving im ...
s of facial sinuses. Its association with allergy has made it a common fungus used in allergen medical testing. Industrial use of the fungus is in the production of enzymes and the manufacture of certain dairy foods.


Morphology and taxonomy

The dimorphic form of the species mainly exists and grows vegetatively as either a filamentous
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
e (mould form) or as spherical
yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constitut ...
(yeast form). However, the organism is best known from the mould form which is characterised by the production of asexual reproductive state consisting of tall (up to 2 cm) needle-like sporangiophores with an apical swelling enclosed by a large
sporangium A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cy ...
filled with ellipsoidal, single-celled, smooth-walled, unpigmented sporangiospores. In the laboratory, the fungus forms dark grey or light grey colonies on most common laboratory media. If subjected to anaerobic conditions, the fungus may convert to the yeast-like form. Anaerobic conditions and 30% carbon dioxide presence stimulate conversion to yeast form. Likewise, cultures supplemented with Tween 80,
ergosterol Ergosterol (ergosta-5,7,22-trien-3β-ol) is a sterol found in cell membranes of fungi and protozoa, serving many of the same functions that cholesterol serves in animal cells. Because many fungi and protozoa cannot survive without ergosterol, the ...
and supplied with 100% nitrogen also converted to yeast. Conversely, increasing oxygen concentration will cause conversion of the yeast form to the mould form. Like many zygomycetes, ''M. racemosus'' reproduces both sexually and asexually depending on environmental conditions. During sexual reproduction, hyphae of compatible mating types touch and fuse, ultimately giving rise to a thick-walled zygosporangium containing a single zygospore. Germination from the zygospore leads to growth of new hyphae that give rise to asexual spores of both + and - mating type. Germination of these spores produces new haploid hyphae of the same mating type.


Physiology and ecology

''M. racemosus'' possesses the ability to exhibit multiple morphology (mainly, filamentous and spherical shape) to withstand various environmental stress. This has given it ability to survive many conditions and it has a worldwide distribution, reported most frequently in Europe as well as Americas. In the tropics, it has been seen at higher altitudes. While the species is primarily soil-based, it has been shown to exist elsewhere such as in horse manure, plant remains, grains, vegetables and nuts. Typically, it is often seen on plant-based materials such as soft fruit, fruit juice and marmalade but it has also been isolated from non-plant sources like soft camembert cheese. ''M. racemosus'' has also been isolated from the human gut
microbiome A microbiome () is the community of microorganisms that can usually be found living together in any given habitat. It was defined more precisely in 1988 by Whipps ''et al.'' as "a characteristic microbial community occupying a reasonably well ...
of non-obese individuals. It is the most common mould found in the floor dust in houses and is largely considered as an indoor mould.''M. racemosus'' is uniquely known for its ability to display multiple morphologies but most of the time, studies are made based on the dimorphic form of the species. It is a
facultative anaerobic A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are '' Staphylococc ...
zygomycote and fast-growing, conferring it ability to survive in multiple conditions/locations all over the world. ''M. racemosus'' possesses the ability to biosynthesize chitin and chitosan, which has been proposed as a mechanism supporting the ability of the fungus to switch between the yeast and the mould phases. Genomic analysis of ''M. racemosus'' has revealed genes similar to human RAS genes, and it is proposed that these genes help with germination and dimorphism.
Protein kinase A In cell biology, protein kinase A (PKA) is a family of enzymes whose activity is dependent on cellular levels of cyclic AMP (cAMP). PKA is also known as cAMP-dependent protein kinase (). PKA has several functions in the cell, including regulatio ...
(PKA) genes such as pkaR are highly also expressed during dimorphic shift.


Human disease

''M. racemosus'' is a rare agent of human disease, typically only associated with opportunistic infection of immunocompromised individuals such as children, elderly and diseased patients (HIV, Ebola etc.). It is an agent of
Mucormycosis Mucormycosis, also known as black fungus, is a serious fungal infection that comes under fulminant fungal sinusitis, usually in people who are immunocompromised. It is curable only when diagnosed early. Symptoms depend on where in the body the ...
, a potentially life-threatening infection often involving the head airways. Pulmonary, cutaneous, and gastrointestinal (GI) infections have also been observed leading to an array of clinical presentations in infected individuals. Risk factors such as
diabetic ketoacidosis Diabetic ketoacidosis (DKA) is a potentially life-threatening complication of diabetes mellitus. Signs and symptoms may include vomiting, abdominal pain, deep gasping breathing, increased urination, weakness, confusion and occasionally loss of ...
and
neutropenia Neutropenia is an abnormally low concentration of neutrophils (a type of white blood cell) in the blood. Neutrophils make up the majority of circulating white blood cells and serve as the primary defense against infections by destroying bacteria ...
are present in most cases. Treatment of ''M. racemosus'' can be difficult due to
histopathologic Histopathology (compound of three Greek words: ''histos'' "tissue", πάθος ''pathos'' "suffering", and -λογία ''-logia'' "study of") refers to the microscopic examination of tissue in order to study the manifestations of disease. Spec ...
differentiation of the fungus. In addition to commonly used antifungal agents, biological compounds like
Lovastatin Lovastatin, sold under the brand name Mevacor among others, is a statin medication, to treat hypercholesterolemia, high blood cholesterol and reduce the risk of cardiovascular disease. Its use is recommended together with lifestyle changes. It ...
, Aleuria aurantia lectin (AAL) and antimicrobial peptides (AMPs LR14) have been isolated and showed antimicrobial effects towards ''M. racemosus''. Allergies to ''M. racemosus'' have been reported to affect immunologically normal individuals from in a range of places (Netherlands, Turkey and Brazil). Allergy to ''M. racemosus'' has been also associated with fungal rhinosinusitis,
rhinitis Rhinitis, also known as coryza, is irritation and inflammation of the mucous membrane inside the nose. Common symptoms are a stuffy nose, runny nose, sneezing, and post-nasal drip. The inflammation is caused by viruses, bacteria, irritants o ...
and
extrinsic allergic alveolitis Hypersensitivity pneumonitis (HP) or extrinsic allergic alveolitis (EAA) is a syndrome caused by the repetitive inhalation of antigens from the environment in susceptible or sensitized people. Common antigens include molds, bacteria, bird dropping ...
.
Asthma Asthma is a long-term inflammatory disease of the airways of the lungs. It is characterized by variable and recurring symptoms, reversible airflow obstruction, and easily triggered bronchospasms. Symptoms include episodes of wheezing, cou ...
tic patients have also shown elevated sensitization to ''M. racemosus''. ''Mucor racemosus''-specific IgE antibody is commonly used and available for medical as well as laboratory use in allergen assay (ImmunoCAP).


Commercial and biotechnological use

The capacity of ''M. racemosus'' to grow as a yeast and its various abilities to manufacture biochemicals have led to its use in industry. For example, it can produce a high yield of
phytase A phytase (''myo''-inositol hexakisphosphate phosphohydrolase) is any type of phosphatase enzyme that catalyzes the hydrolysis of phytic acid (myo-inositol hexakisphosphate) – an indigestible, organic form of phosphorus that is found in many pl ...
, an important
industrial enzyme Industrial enzymes are enzymes that are commercially used in a variety of industries such as Pharmaceutical drug, pharmaceuticals, chemical production, biofuels, food processing, food & beverage, and consumer products. Due to advancements in recent ...
. It also has an increased extracellular
protease A protease (also called a peptidase, proteinase, or proteolytic enzyme) is an enzyme that catalyzes (increases reaction rate or "speeds up") proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the ...
activity, suggesting its biotechnological suitability for the production of other
industrial enzymes Industrial enzymes are enzymes that are commercially used in a variety of industries such as pharmaceuticals, chemical production, biofuels, food & beverage, and consumer products. Due to advancements in recent years, biocatalysis through isolat ...
. It is also used in production of
homeopathic Homeopathy or homoeopathy is a pseudoscientific system of alternative medicine. It was conceived in 1796 by the German physician Samuel Hahnemann. Its practitioners, called homeopaths, believe that a substance that causes symptoms of a dise ...
medicinal products. In the manufacture of sufu (fermented cheese-like soybean product common in China and Vietnam), the fungal fermentation of soybean curd (tofu) results in moulded tofu, ''pehtze''. The final product (sufu) is obtained by maturing ''pehtze'' in a brine containing alcohol and salt for several months. It possesses the ability to adapt phenotypically to several different
antibiotics An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention o ...
after exposure to a single drug, which makes it a good model for phenotypic multidrug resistance in lower
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s. It has been shown to adapt to famous antibiotics like
cycloheximide Cycloheximide is a naturally occurring fungicide produced by the bacterium ''Streptomyces griseus''. Cycloheximide exerts its effects by interfering with the translocation step in protein synthesis (movement of two tRNA molecules and mRNA in rela ...
, trichodermin and
amphotericin B Amphotericin B is an antifungal medication used for serious fungal infections and leishmaniasis. The fungal infections it is used to treat include mucormycosis, aspergillosis, blastomycosis, candidiasis, coccidioidomycosis, and cryptococcosis. Fo ...
. Cells adapted to cycloheximide particularly have been observed to be 40-times more resistant than non-adapted cells to the drug. These adapted cells have been studied to better understand their greater efficiency of membrane transport (efflux of drugs). ''Mucor racemosus'' can biotransform lipids like 4-ene-3-one steroids and 20(S)-Protopanaxatriol into several different products, some of which have
anticancer An anticarcinogen (also known as a carcinopreventive agent) is a substance that counteracts the effects of a carcinogen or inhibits the development of cancer. Anticarcinogens are different from anticarcinoma agents (also known as anticancer or an ...
properties (as the metabolites resulted in increased intracellular calcium ion content, leading to cell cycle arrest and
apoptosis Apoptosis (from grc, ἀπόπτωσις, apóptōsis, 'falling off') is a form of programmed cell death that occurs in multicellular organisms. Biochemical events lead to characteristic cell changes (morphology) and death. These changes incl ...
). Two of the products formed from this
biotransformation Biotransformation is the biochemical modification of one chemical compound or a mixture of chemical compounds. Biotransformations can be conducted with whole cells, their lysates, or purified enzymes. Increasingly, biotransformations are effected w ...
are two novel hydroperoxylated metabolites that have been shown to be effective against
prostate cancer Prostate cancer is cancer of the prostate. Prostate cancer is the second most common cancerous tumor worldwide and is the fifth leading cause of cancer-related mortality among men. The prostate is a gland in the male reproductive system that sur ...
cells. Secondary metabolites of ''M. racemosus'' do not exhibit genotoxic activity, and the species is not known to be a producer of
mycotoxin A mycotoxin (from the Greek μύκης , "fungus" and τοξίνη , "toxin") is a toxic secondary metabolite produced by organisms of kingdom Fungi and is capable of causing disease and death in both humans and other animals. The term 'mycotoxin' ...
s. However, some secondary metabolites of the fungus have been found to have anti-inflammatory activity similar to the drug
dexamethasone Dexamethasone is a glucocorticoid medication used to treat rheumatic problems, a number of skin diseases, severe allergies, asthma, chronic obstructive lung disease, croup, brain swelling, eye pain following eye surgery, superior vena cav ...
.


References

{{Taxonbar, from=Q2450163 Mucoraceae Fungi described in 1791