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The mycobiome, mycobiota, or fungal
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 wel ...
, is the fungal community in and on an organism. The word “mycobiome” comes from the ancient Greek μύκης (mukēs), meaning "fungus" with the suffix “biome” derived from the Greek βίος (bíos), meaning “life.” The term was first coined in the 2009 paper by Gillevet et al. Most species of fungi are decomposers with the ability to break down complex polymers. Fungi are commonly found within plant cells in an
endophytic An endophyte is an endosymbiont, often a bacterium or fungus, that lives within a plant for at least part of its life cycle without causing apparent disease. Endophytes are ubiquitous and have been found in all species of plants studied to date; h ...
relationship or as a
pathogen In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a g ...
. Most plants also form mutualistic relationships with fungi that accelerate nutrient uptake among their root structures. The most common phyla present in the fungal communities that live alongside animals and in aquatic environments are
Ascomycota 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 defi ...
and
Basidiomycota 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. More specifically, Bas ...
. Animals will typically form a
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit fro ...
relationship with fungi with the occasional occurrence of a pathogenic interaction.


Interactions with other microbes

Fungal microbes are amongst a wide variety of other microbes involved in a symbiotic relationship involving multicellular organisms. In mammals, the
gut flora Gut microbiota, gut microbiome, or gut flora, are the microorganisms, including bacteria, archaea, fungi, and viruses that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the gut m ...
is usually met with vastly diverse populations of microbes from many kingdoms, where fungal populations make up less than 1% of the entire gut biome. Due to the coexistence of fungal populations with other microbes in most cases of host-symbiont associations, it’s important to assess common dynamics that may occur. Most interactions between microbes in the gut are either
competitive Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indivi ...
or cooperative. This can be seen with multiple fungal microbes as well by observing populations through the treatment of
antibiotic 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 ...
s and
antifungal An antifungal medication, also known as an antimycotic medication, is a pharmaceutical fungicide or fungistatic used to treat and prevent mycosis such as athlete's foot, ringworm, candidiasis (thrush), serious systemic infections such as ...
s. Research on microbial populations in animal models has resulted in noticeable fluctuations in microbe populations. Antibiotic treatment has mostly shown increases in parasitic fungal presence, suggesting competitive behaviors between microbes against fungi. Additionally, application of antifungal molecules have resulted in
colitis Colitis is swelling or inflammation of the large intestine ( colon). Colitis may be acute and self-limited or long-term. It broadly fits into the category of digestive diseases. In a medical context, the label ''colitis'' (without qualification ...
in mice, suggesting that commensal fungi are responsible for balancing bacterial populations that are involved in
dysbiosis Dysbiosis (also called dysbacteriosis) is characterized by a disruption to the microbiome resulting in an imbalance in the microbiota, changes in their functional composition and metabolic activities, or a shift in their local distribution. For e ...
. Despite limited knowledge on the gut mycobiome, this research suggests that interactions between fungal and bacterial microbes in the mammalian gut are largely competitive.


Lichen

Lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.photosynthesizing Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
microbes that are either algal chlorophytes or
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, bl ...
. The primary symbiosis involves fungi gaining energy from their photosynthesizing symbiotic microbe, and in return, provides a structural scaffold from the hyphae. These interactions are classically seen as mutualistic, but have been questioned if fungi might also be capable of playing a controlled parasitic role.


Plants

Plants also have mutualistic
symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or para ...
relationships with fungal communities that are found in a microbe abundant layer of the soil called the
rhizosphere The rhizosphere is the narrow region of soil or substrate that is directly influenced by root secretions and associated soil microorganisms known as the root microbiome. Soil pores in the rhizosphere can contain many bacteria and other microo ...
. Fungi can be vertically transmitted to progeny plants, or horizontally through fungal diffusion in the soil. Regardless of transmission, the most common cases of fungal plant symbiosis happens when fungal communities colonize plant root structure. There are some cases of symbiosis that Begin before maturity such as the
Orchidaceae Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Along with the Asteraceae, they are one of the two largest families of flowerin ...
family, in which symbiosis begins at the seed
germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fe ...
phase. Arbuscular
mycorrhiza   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plan ...
l fungi supply the plant essential inorganic nutrients (in the form of minerals) for 80% of terrestrial plant species. In return the plant will provide fungi with plant assimilated carbon that can easily be metabolized and used for energy.


Evolutionary Symbiosis

DNA sequencing and fossil records have derived evidence that this mutualistic relationship between fungi and plants is extremely ancient. It is even suggested that different subspecies of fungi have played major roles in shaping the plants immune system in the early evolutionary moments of plants colonizing land. Specific
gene cluster A gene family is a set of homologous genes within one organism. A gene cluster is a group of two or more genes found within an organism's DNA that encode similar polypeptides, or proteins, which collectively share a generalized function and are ...
s in the plant's
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
used to code for
fungicidal Fungicides are biocidal chemical compounds or biological organisms used to kill parasitic fungi or their spores. A fungistatic inhibits their growth. Fungi can cause serious damage in agriculture, resulting in critical losses of yield, quality ...
properties like the production of
isoflavonoid Isoflavonoids are a class of flavonoid phenolic compounds, many of which are biologically active. Isoflavonoids and their derivatives are sometimes referred to as phytoestrogens, as many isoflavonoid compounds have biological effects via the estr ...
s in the
Fabaceae The Fabaceae or Leguminosae,International Code of Nomenc ...
plant family are toxic and used to regulate the growth of expected fungal presence. In a
coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
ary response, arbuscular fungal communities breed quickly and evolutionarily selected for the development of specific gene clusters to detoxify and metabolize a wide variety of organic antifungal chemicals. This exchange of evolutionary pressures has given a variety of plant species better immune systems, while simultaneously giving mutualist fungal communities new sources of plant energy at the genomic level.


Insects

Interactions between fungi and insects are incredibly common and most of these relationships are either commensal or pathogenic.


Commensal

The commensal relationships typically benefit the insect by allowing them to digest tough materials such as wood. For example, wood-ingesting passalid beetle guts are populated with
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
,
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 constit ...
s, and other
eukaryotic 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 ...
organisms that aid in digestion and nutrient absorption. Without the help of the yeasts to break down
xylose Xylose ( grc, ξύλον, , "wood") is a sugar first isolated from wood, and named for it. Xylose is classified as a monosaccharide of the aldopentose type, which means that it contains five carbon atoms and includes an aldehyde functional g ...
from plant cells, these beetles would be unable to efficiently digest this tough material.


Pathogenic

Insect pathogenic fungi slowly kill their hosts while replicating rapidly inside the infected insect. They typically attach to the external surface of the insect as
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s before germinating, perforating the
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
, and colonizing the internal cavity. The insect is typically killed and new spores are dispersed from the reproductive structures in the body of the insect. A prominent example of an entomopathogenic fungus is
Ophiocordyceps unilateralis ''Ophiocordyceps unilateralis'', commonly known as zombie-ant fungus, is an insect-pathogenic fungus, discovered by the British naturalist Alfred Russel Wallace in 1859, and currently found predominantly in tropical forest ecosystems. ''O. unil ...
, nicknamed the “zombie-ant” fungus. The fungus infects ants and alters their natural behavioral patterns causing the ant to leave their usual environment in the trees in favor of the forest floor – a more suitable environment for fungal growth. The insects will then attach themselves to the underside of a leaf until it dies. Fungal spores then sprout out of the dead ant’s head and are dispersed. One of the first descriptions of insect pathogenic fungi was published in 1835 during an infestation of
Beauveria bassiana ''Beauveria bassiana'' is a fungus that grows naturally in soils throughout the world and acts as a parasite on various arthropod species, causing white muscardine disease; it thus belongs to the entomopathogenic fungi. It is used as a biologic ...
in a silkworm population that upset the silk industry.


Humans

There is a low abundance of fungi associated with most human body sites, such as the gastrointestinal tract, where fungi typically compose just 0.001 - 0.1% of the microbial community. However, fungi compose a significant fraction of the microbiome at some locations, such as the ear canal. The mycobiome is relevant to human physiology as fungi may be important in maintaining microbial community structure, metabolic function, and immune-priming. Mutualism between humans and fungi is not yet well understood, and there is much to be learned about how fungi interact with the nonfungal constituents of the microbiome. Many human diseases, including
hepatitis Hepatitis is inflammation of the liver tissue. Some people or animals with hepatitis have no symptoms, whereas others develop yellow discoloration of the skin and whites of the eyes ( jaundice), poor appetite, vomiting, tiredness, abdominal ...
,
cystic fibrosis Cystic fibrosis (CF) is a rare genetic disorder that affects mostly the lungs, but also the pancreas, liver, kidneys, and intestine. Long-term issues include difficulty breathing and coughing up mucus as a result of frequent lung infections. Ot ...
,
obesity Obesity is a medical condition, sometimes considered a disease, in which excess body fat has accumulated to such an extent that it may negatively affect health. People are classified as obese when their body mass index (BMI)—a person's ...
, and
inflammatory bowel disease Inflammatory bowel disease (IBD) is a group of inflammatory conditions of the colon and small intestine, Crohn's disease and ulcerative colitis being the principal types. Crohn's disease affects the small intestine and large intestine, as well ...
have been associated with changes in the human mycobiome, and it has been proposed that any fungal colonization of the GI tract is a sign of disease. Moreover, detecting tumor-associated mycobiomes may be a novel way of
cancer screening Cancer screening aims to detect cancer before symptoms appear. This may involve blood tests, urine tests, DNA tests, other tests, or medical imaging. The benefits of screening in terms of cancer prevention, early detection and subsequent treatment ...
– it could be used in synergy with other
biomarker In biomedical contexts, a biomarker, or biological marker, is a measurable indicator of some biological state or condition. Biomarkers are often measured and evaluated using blood, urine, or soft tissues to examine normal biological processes, p ...
s such as of bacteriomes. Fungal interactions in the human host can be opportunistic, mutualistic, parasitic, commensalistic, and amensalistic. Though there is a small amount of fungi considered to be true pathogens due to causing disease in healthy individuals, the majority of fungi are considered opportunistic, only affecting an immunosuppressed host. In these cases,  pathogen-mediated damage is dependent upon the host; disease could only occur if  the host has either a hyperactive or  weakened immune system. Commensal relationships between human host and pathogen are possible  when the fungi are present in a host for long periods of time without causing disease. One example of a fungus that forms a commensal relationship in a human host is Cryptococcus. Candida albicans can also form a long-time commensal relationship with its host. Some fungi are niche specific symbionts in the human host. Some fungi can be introduced to the host from the environment by skin contact, oral or respiratory routes, while others are acquired vertically through birth. In the human GI tract, there are about 50 observed genera of fungi. Some are acquired through what the host eats; as such showing that the stability of the fungal population is dependent on the diet of the host and host immune status. About 20 genera of fungi have been observed in the vaginal niche. Most fungi that colonize the vagina show a commensalistic relationship with the host, but factors like antibiotic uses and pregnancy could influence the vaginal mycobiome.


References

{{reflist Eukaryotic microbiology Fungus ecology Microbiomes