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''Candida'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom (biology), kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are est ...
s. It is the most common cause of fungal infections worldwide and the largest genus of medically important yeasts. The genus ''Candida'' encompasses about 200 species. Many species are harmless commensals or
endosymbiont An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualism (biology), mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), whi ...
s of hosts including
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s. When mucosal barriers are disrupted or the
immune system The immune system is a network of biological systems that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to bacteria, as well as Tumor immunology, cancer cells, Parasitic worm, parasitic ...
is compromised, however, they can invade and cause disease, known as an
opportunistic infection An opportunistic infection is an infection that occurs most commonly in individuals with an immunodeficiency disorder and acts more severe on those with a weakened immune system. These types of infections are considered serious and can be caused b ...
. Candida is located on most mucosal surfaces and mainly the
gastrointestinal tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the Digestion, digestive system that leads from the mouth to the anus. The tract is the largest of the body's systems, after the cardiovascula ...
, along with the skin. '' Candida albicans'' is one of the most commonly isolated species and can cause infections ( candidiasis or thrush) in humans and other animals. In
winemaking Winemaking, wine-making, or vinification is the production of wine, starting with the selection of the fruit, its Ethanol fermentation, fermentation into alcohol, and the bottling of the finished liquid. The history of wine-making stretches over ...
, some species of ''Candida'' can potentially spoil wines. Many species are found in
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 g ...
, including ''C. albicans'' in
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
ian hosts, whereas others live as endosymbionts in insects. Systemic infections of the bloodstream and major organs ( candidemia or invasive candidiasis), particularly in patients with an impaired immune system (
immunocompromised Immunodeficiency, also known as immunocompromise, is a state in which the immune system's ability to fight infectious diseases and cancer is compromised or entirely absent. Most cases are acquired ("secondary") due to extrinsic factors that affe ...
), affect over 90,000 people a year in the US. The
genome 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 genes, other functional regions of the genome such as ...
of several ''Candida'' species has been sequenced.
Antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting pathogenic bacteria, bacterial infections, and antibiotic medications are widely used in the therapy ...
s promote yeast (fungal) infections, including gastrointestinal (GI) ''Candida'' overgrowth and penetration of the GI mucosa. While women are more susceptible to genital yeast infections, men can also be infected. Certain factors, such as prolonged antibiotic use, increase the risk for both men and women. People with
diabetes Diabetes mellitus, commonly known as diabetes, is a group of common endocrine diseases characterized by sustained high blood sugar levels. Diabetes is due to either the pancreas not producing enough of the hormone insulin, or the cells of th ...
or the immunocompromised, such as those infected with HIV, are more susceptible to yeast infections. '' Candida antarctica'' and '' Candida rugosa'' are a source of industrially important
lipase In biochemistry, lipase ( ) refers to a class of enzymes that catalyzes the hydrolysis of fats. Some lipases display broad substrate scope including esters of cholesterol, phospholipids, and of lipid-soluble vitamins and sphingomyelinases; howe ...
s, while '' Candida krusei'' is prominently used to ferment cacao during chocolate production. Lipases from '' Candida rugosa'' are also used to digest
fat In nutrition science, nutrition, biology, and chemistry, fat usually means any ester of fatty acids, or a mixture of such chemical compound, compounds, most commonly those that occur in living beings or in food. The term often refers specif ...
s in laboratory assays because of their broad range of activity.


Biology

When grown in a
laboratory A laboratory (; ; colloquially lab) is a facility that provides controlled conditions in which scientific or technological research, experiments, and measurement may be performed. Laboratories are found in a variety of settings such as schools ...
, ''Candida'' appears as large, round, white or cream (''albicans'' means "whitish" in
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
) colonies, which emit a yeasty odor on
agar plate An agar plate is a Petri dish that contains a growth medium solidified with agar, used to Microbiological culture, culture microorganisms. Sometimes selective compounds are added to influence growth, such as antibiotics. Individual microorganism ...
s at room temperature. ''C. albicans'' ferments
glucose Glucose is a sugar with the Chemical formula#Molecular formula, molecular formula , which is often abbreviated as Glc. It is overall the most abundant monosaccharide, a subcategory of carbohydrates. It is mainly made by plants and most algae d ...
and
maltose } Maltose ( or ), also known as maltobiose or malt sugar, is a disaccharide formed from two units of glucose joined with an α(1→4) bond. In the isomer isomaltose, the two glucose molecules are joined with an α(1→6) bond. Maltose is the tw ...
to acid and gas,
sucrose Sucrose, a disaccharide, is a sugar composed of glucose and fructose subunits. It is produced naturally in plants and is the main constituent of white sugar. It has the molecular formula . For human consumption, sucrose is extracted and refined ...
to acid, and does not ferment
lactose Lactose is a disaccharide composed of galactose and glucose and has the molecular formula C12H22O11. Lactose makes up around 2–8% of milk (by mass). The name comes from (Genitive case, gen. ), the Latin word for milk, plus the suffix ''-o ...
, which helps to distinguish it from other ''Candida'' species. Recent molecular
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
studies show that the genus ''Candida'', as currently defined, is extremely
polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
(encompassing distantly-related species that do not form a natural group). Before the advent of inexpensive molecular methods,
yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom (biology), kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are est ...
s that were isolated from infected patients were often called ''Candida'' without clear evidence of relationship to other ''Candida'' species. For example, '' Candida glabrata'', '' Candida guilliermondii'', and '' Candida lusitaniae'' are clearly misclassified and will be placed in other
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
once phylogenetic reorganization is complete (for example, see Khunnamwong ''et al.'' 2015). Some species of ''Candida'' use a non-standard
genetic code Genetic code is a set of rules used by living cell (biology), cells to Translation (biology), translate information encoded within genetic material (DNA or RNA sequences of nucleotide triplets or codons) into proteins. Translation is accomplished ...
in the
translation Translation is the communication of the semantics, meaning of a #Source and target languages, source-language text by means of an Dynamic and formal equivalence, equivalent #Source and target languages, target-language text. The English la ...
of their nuclear genes into the
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the 22 α-amino acids incorporated into proteins. Only these 22 a ...
sequences of polypeptides. The difference in the genetic code between species possessing this alternative code is that the codon CUG (normally encoding the amino acid
leucine Leucine (symbol Leu or L) is an essential amino acid that is used in the biosynthesis of proteins. Leucine is an α-amino acid, meaning it contains an α-amino group (which is in the protonated −NH3+ form under biological conditions), an α-Car ...
) is translated by the yeast as a different amino acid, serine. The alternative translation of the CUG codon in these species is due to a novel nucleic acid sequence in the serine-
tRNA Transfer ribonucleic acid (tRNA), formerly referred to as soluble ribonucleic acid (sRNA), is an adaptor molecule composed of RNA, typically 76 to 90 nucleotides in length (in eukaryotes). In a cell, it provides the physical link between the gene ...
(ser-tRNACAG), which has a guanosine located at position 33, 5' to the anticodon. In all other tRNAs, this position is normally occupied by a
pyrimidine Pyrimidine (; ) is an aromatic, heterocyclic, organic compound similar to pyridine (). One of the three diazines (six-membered heterocyclics with two nitrogen atoms in the ring), it has nitrogen atoms at positions 1 and 3 in the ring. The oth ...
(often
uridine Uridine (symbol U or Urd) is a glycosylated pyrimidine analog containing uracil attached to a ribose ring (or more specifically, a ribofuranose) via a β-N1- glycosidic bond. The analog is one of the five standard nucleosides which make up nuc ...
). This genetic code change is the only such known alteration in cytoplasmic
mRNA In molecular biology, messenger ribonucleic acid (mRNA) is a single-stranded molecule of RNA that corresponds to the genetic sequence of a gene, and is read by a ribosome in the process of Protein biosynthesis, synthesizing a protein. mRNA is ...
, in both the
prokaryote A prokaryote (; less commonly spelled procaryote) is a unicellular organism, single-celled organism whose cell (biology), cell lacks a cell nucleus, nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Ancient Gree ...
s, and the
eukaryote The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...
s, involving the reassignment of a sense codon. This novel genetic code may be a mechanism for more rapid adaptation to the organism's environment, as well as playing an important role in the
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
of the genus ''Candida'' by creating genetic barriers that encouraged
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
.


Pathogen

''Candida'' are almost universal in low numbers on healthy adult
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other animal coverings, such as the arthropod exoskeleton, have different ...
and ''C. albicans'' is part of the normal flora of the mucous membranes of the respiratory, gastrointestinal and female genital tracts. The dryness of skin compared to other tissues prevents the growth of the fungus, but damaged skin or skin in intertriginous regions is more amenable to rapid growth. A severe consequence of candida infection can be esophageal cancer, overgrowth with candida in the
esophagus The esophagus (American English), oesophagus (British English), or œsophagus (Œ, archaic spelling) (American and British English spelling differences#ae and oe, see spelling difference) all ; : ((o)e)(œ)sophagi or ((o)e)(œ)sophaguses), c ...
is the most important risk factor for developing esophageal cancer in patients with
achalasia Esophageal achalasia, often referred to simply as achalasia, is a failure of smooth muscle fibers to relax, which can cause the lower esophageal sphincter to remain closed. Without a modifier, "achalasia" usually refers to achalasia of the esopha ...
. Overgrowth of several species, including ''C. albicans'', can cause infections ranging from superficial, such as oropharyngeal candidiasis (thrush) or vulvovaginal candidiasis (vaginal candidiasis) and subpreputial candidiasis, which may cause
balanitis Balanitis is inflammation of the glans penis. Balanoposthitis is the proper term when the foreskin is also affected. Balanitis on boys in diapers must be distinguished from redness caused by ammoniacal dermatitis. Etymology The word ''balanit ...
, to systemic, such as fungemia and invasive candidiasis. Oral candidiasis is common in elderly denture-wearers. In otherwise healthy individuals, these superficial infections can be cured with topical or systemic antifungal medications (commonly over-the-counter antifungal treatments like miconazole or
clotrimazole Clotrimazole, sold under the brand name Lotrimin, among others, is an antifungal medication. It is used to treat vaginal yeast infections, oral thrush, diaper rash, tinea versicolor, and types of ringworm including athlete's foot and jock itc ...
). In debilitated or immunocompromised patients, or if introduced intravenously (into the bloodstream), candidiasis may become a systemic disease producing
abscess An abscess is a collection of pus that has built up within the tissue of the body, usually caused by bacterial infection. Signs and symptoms of abscesses include redness, pain, warmth, and swelling. The swelling may feel fluid-filled when pre ...
es,
thrombophlebitis Thrombophlebitis is a phlebitis (inflammation of a vein) related to a thrombus (blood clot). When it occurs repeatedly in different locations, it is known as thrombophlebitis migrans (migratory thrombophlebitis). Signs and symptoms The following ...
, endocarditis, or infections of the eyes or other organs. Typically, relatively severe neutropenia (low
neutrophil Neutrophils are a type of phagocytic white blood cell and part of innate immunity. More specifically, they form the most abundant type of granulocytes and make up 40% to 70% of all white blood cells in humans. Their functions vary in differe ...
s) is a prerequisite for ''Candida'' to pass through the defenses of the skin and cause disease in deeper tissues; in such cases, mechanical disruption of the infected skin sites is typically a factor in the fungal invasion of the deeper tissues. The most common way to treat invasive candida infections is with the use of amphotericin or fluconazole; other methods would include surgery.


Applications

''C. albicans'' has been used in combination with
carbon nanotubes A carbon nanotube (CNT) is a tube made of carbon with a diameter in the nanometre range (nanoscale). They are one of the allotropes of carbon. Two broad classes of carbon nanotubes are recognized: * ''Single-walled carbon nanotubes'' (''SWC ...
(CNT) to produce stable electrically conductive bio-nano-composite tissue materials that have been used as temperature-sensing elements.


Species

Among ''Candida'' species, ''C. albicans'', which is a normal constituent of the human flora, a commensal of the skin and the gastrointestinal and genitourinary tracts, is responsible for the majority of ''Candida'' bloodstream infections (candidemia). Yet, there is an increasing incidence of infections caused by '' C. glabrata'' and '' C. rugosa'', which could be because they are frequently less susceptible to the currently used azole-group of antifungals. Other medically important species include '' C. parapsilosis'', '' C. tropicalis'', '' C. dubliniensis''. and the more recently emerging pathogen '' C. auris''. Other ''Candida'' species, such as '' C. oleophila'', have been used as
biological control Biological control or biocontrol is a method of controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or o ...
agents in fruit. * '' C. albicans'' * '' C. ascalaphidarum'' * '' C. amphixiae'' * '' C. antarctica'' * '' C. argentea'' * '' C. atlantica'' * '' C. atmosphaerica'' * '' C. auris'' * '' C. blankii'' * '' C. blattae'' * '' C. boidinii'' * '' C. bracarensis'' * '' C. bromeliacearum'' * '' C. carpophila'' * '' C. carvajalis'' * '' C. catenulata'' * '' C. cerambycidarum'' * '' C. chauliodes'' * '' C. corydali'' * '' C. crusei'' * '' C. dosseyi'' * '' C. dubliniensis'' * '' C. ergatensis'' * '' C. fructus'' * '' C. glabrata'' * '' C. fermentati'' * '' C. guilliermondii'' * '' C. haemulonii'' * '' C. humilis'' * '' C. insectamens'' * '' C. insectorum'' * '' C. intermedia'' * '' C. jeffresii'' * '' C. kefyr'' * '' C. keroseneae'' * '' C. krusei'' * '' C. lipolytica'' * '' C. lusitaniae'' * '' C. lyxosophila'' * '' C. maltosa'' * '' C. marina'' * '' C. membranifaciens'' * '' C. mogii'' * '' C. oleophila'' * '' C. oregonensis'' * '' C. parapsilosis'' * '' C. quercitrusa'' * '' C. rhizophoriensis'' * '' C. rugosa'' * '' C. sake'' * '' C. sharkiensis'' * '' C. shehatea'' * '' C. temnochilae'' * '' C. tenuis'' * '' C. theae'' * '' C. tolerans'' * '' C. tropicalis'' * '' C. tsuchiyae'' * '' C. sinolaborantium'' * '' C. sojae'' * '' C. subhashii'' * '' C. viswanathii'' * '' C. ubatubensis'' * '' C. utilis'' * '' C. zemplinina''


References


External links


Labs working on Candida
* {{Authority control Yeasts Gut flora Pathogenic microbes