A heterokaryon is a

multinucleate Multinucleate cells (also known as multinucleated or polynuclear cells) are eukaryotic cells that have more than one nucleus per cell, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordinat ...

cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...

that contains genetically different nuclei. Heterokaryotic and heterokaryosis are derived terms. This is a special type of

syncytium A syncytium (; plural syncytia; from Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") or symplasm is a multinucleate cell which can result from multiple cell fusions of uninuclear cells (i.e., cells with a single nucleus) ...

. This can occur naturally, such as in the

mycelium Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrate (biology), substrates. A typical single ...

fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...

during sexual reproduction, or artificially as formed by the experimental fusion of two genetically different cells, as e.g., in hybridoma technology.

Etymology

''Heterokaryon'' is from neo-classic

Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...

''hetero'', meaning ''different'', and ''karyon'', meaning ''kernel'' or in this case ''nucleus''. The term was coined in 1965, independently by B. Ephrussi and M. Weiss, by H. Harris and J. F. Watkins, and by Y. Okada and F. Murayama.

Occurrence

Heterokaryons are found in the life cycle of yeasts, for example ''Saccharomyces cerevisiae'', a genetic model organism. The heterokaryon stage is produced from the fusion of two haploid cells. This ''transient'' heterokaryon can produce further haploid buds, or cell nuclei can fuse and produce a diploid cell, which can then undergo mitosis.

Ciliate protozoans

The term was first used for

ciliate The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to flagellum, eukaryotic flagella, but are in general shorter and present in much larger numbers, with a ...

protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...

ns such as ''

Tetrahymena ''Tetrahymena'', a Unicellular organism, unicellular eukaryote, is a genus of free-living ciliates. The genus Tetrahymena is the most widely studied member of its phylum. It can produce, store and react with different types of hormones. Tetrah ...

''. This has two types of cell nuclei, a large,

somatic Somatic may refer to: * Somatic (biology), referring to the cells of the body in contrast to the germ line cells ** Somatic cell, a non-gametic cell in a multicellular organism * Somatic nervous system, the portion of the vertebrate nervous sys ...

macronucleus A macronucleus (formerly also meganucleus) is the larger type of nucleus in ciliates. Macronuclei are polyploid and undergo direct division without mitosis. It controls the non-reproductive cell functions, such as metabolism. During conjugation, ...

and a small,

germline In biology and genetics, the germline is the population of a multicellular organism's cells that pass on their genetic material to the progeny (offspring). In other words, they are the cells that form the egg, sperm and the fertilised egg. They ...

micronucleus Micronucleus is the name given to the small nucleus that forms whenever a chromosome or a fragment of a chromosome is not incorporated into one of the daughter nuclei during cell division. It usually is a sign of genotoxic events and chromosomal i ...

. Both exist in a single cell at the same time and carry out different functions with distinct

cytological Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living and ...

and

biochemical Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology an ...

properties.

True fungi

Many fungi (notably the

arbuscular mycorrhizal fungi An arbuscular mycorrhiza (AM) (plural ''mycorrhizae'', a.k.a. ''endomycorrhiza'') is a type of mycorrhiza in which the symbiont fungus (''AM fungi'', or AMF) penetrates the cortical cells of the roots of a vascular plant forming arbuscules. (N ...

) exhibit heterokaryosis. The

haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...

nuclei within a mycelium may differ from one another not merely by accumulating

mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mi ...

s, but by the non-sexual fusion of genetically distinct fungal

hyphae 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 ...

, although a self / non-self recognition system exists in Fungi and usually prevents fusions with non-self. Heterokaryosis is also common upon mating, as in Dikarya (

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 def ...

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, Basi ...

). Mating requires the encounter of two haploid nuclei of compatible mating types. These nuclei do not immediately fuse, and remain haploid in a n+n state until the very onset of meiosis: this phenomenon is called delayed karyogamy. Heterokaryosis can lead to individuals that have different nuclei in different parts of their mycelium, although in ascomycetes, particularly in "

Neurospora ''Neurospora'' is a genus of Ascomycete fungi. The genus name, meaning "nerve spore" refers to the characteristic striations on the spores that resemble axons. The best known species in this genus is '' Neurospora crassa'', a common model organ ...

", nuclei have been shown to flow and mix throughout the mycelium. In heterokaryons, the notion of ''individual'' itself becomes vague since the rule of “one genome = one individual” does not apply any more. Genetic heterogeneity within an individual is indeed usually considered to be detrimental, as selfish variants may be selected for and disrupt the integrity of the individual level.

Slime molds

Heterokaryosis is most common in fungi, but also occurs in

slime mold Slime mold or slime mould is an informal name given to several kinds of unrelated eukaryotic organisms with a life cycle that includes a free-living single-celled stage and the formation of spores. Spores are often produced in macroscopic mu ...

s. This happens because the nuclei in the 'plasmodium' form are the products of many pairwise fusions between

amoeboid An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudopo ...

individuals. When genetically divergent nuclei come together in the plasmodium form, cheaters have been shown to emerge. However, genetic homogeneity among fusing amoeboid serves to maintain the multicellular plasmodium.Kuzdzal-Fick, J. J., S. A. Fox, J. E. Strassmann, and D. C. Queller. 2011. High Relatedness is Necessary and Sufficient to Maintain Multicellularity in Dictyostelium. ''Science'' 334:1548-1551

Artificial heterokaryon

A medical example is a heterokaryon composed of nuclei from

Hurler syndrome Hurler syndrome, also known as mucopolysaccharidosis Type IH (MPS-IH), Hurler's disease, and formerly gargoylism, is a genetic disorder that results in the buildup of large sugar molecules called glycosaminoglycans (GAGs) in lysosomes. The inab ...

and

Hunter syndrome Hunter syndrome, or mucopolysaccharidosis type II (MPS II), is a rare genetic disorder in which large sugar molecules called glycosaminoglycans (or GAGs or mucopolysaccharides) build up in body tissues. It is a form of lysosomal storage disease. ...

. Both of these diseases result in problems in mucopolysaccharide metabolism. However, a heterokaryon of nuclei from both of these diseases exhibits normal mucopolysaccharide metabolism, proving that the two syndromes affect different proteins and so can correct each other in the heterokaryon.

References

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External links

MedicineNet.com
Eukaryotic cells Mycology Cell biology