Heterokaryosis
Encyclopedia
Heterokaryosis is a term used in biology
Biology
Biology is a natural science concerned with the study of life and living organisms, including their structure, function, growth, origin, evolution, distribution, and taxonomy. Biology is a vast subject containing many subdivisions, topics, and disciplines...

 meaning to have two or more genetically different nuclei within the same mycelium
Mycelium
thumb|right|Fungal myceliaMycelium is the vegetative part of a fungus, consisting of a mass of branching, thread-like hyphae. The mass of hyphae is sometimes called shiro, especially within the fairy ring fungi. Fungal colonies composed of mycelia are found in soil and on or within many other...

 of a fungus
Fungus
A fungus is a member of a large 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, Fungi, which is separate from plants, animals, and bacteria...

 or other life form. This is a special type of syncytium
Syncytium
In biology, a syncytium is a large cell-like structure; filled with cytoplasm and containing many nuclei. Most cells in eukaryotic organisms have a single nucleus; syncytia are specialized forms used by various organisms.The term may also refer to cells that are connected by specialized membrane...

.

A heterokaryon is a cell with more than one nucleus of differing genetic origin. The term was first used for ciliate
Ciliate
The ciliates are a group of protozoans characterized by the presence of hair-like organelles called cilia, which are identical in structure to flagella but typically shorter and present in much larger numbers with a different undulating pattern than flagella...

 protozoa
Protozoa
Protozoa are a diverse group of single-cells eukaryotic organisms, many of which are motile. Throughout history, protozoa have been defined as single-cell protists with animal-like behavior, e.g., movement...

ns such as Tetrahymena
Tetrahymena
Tetrahymena are free-living ciliate protozoa that can also switch from commensalistic to pathogenic modes of survival. They are common in fresh-water. Tetrahymena species used as model organisms in biomedical research are T. thermophila and T. pyriformis.- T...

. This has two types of cell nuclei, a large, somatic
Somatic
The term somatic means 'of the body',, relating to the body. In medicine, somatic illness is bodily, not mental, illness. The term is often used in biology to refer to the cells of the body in contrast to the germ line cells which usually give rise to the gametes...

 macronucleus
Macronucleus
A macronucleus is the larger type of nucleus in ciliates. Macronuclei are polyploid and undergo direct division without mitosis. It controls the non-reproductive cell functions, the everyday tasks, such as metabolism...

 and a small, germline
Germline
In biology and genetics, the germline of a mature or developing individual is the line of germ cells that have genetic material that may be passed to a child.For example, gametes such as the sperm or the egg, are part of the germline...

 micronucleus
Micronucleus
A the micronucleus is the smaller nucleus in ciliate protozoans, such as the paramecium. In fission it divides by mitosis, and in conjugation furnishes the pairing of gamete nuclei, by whose reciprocal fusion a zygote nucleus is formed, which gives rise to the macronuclei and micronuclei of the...

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

Many fungi (notably the arbuscular mycorrhizal fungi
Mycorrhiza
A mycorrhiza is a symbiotic association between a fungus and the roots of a vascular plant....

) exhibit heterokaryosis. The haploid nuclei within a mycelium may differ from one another not merely by accumulating mutation
Mutation
In molecular biology and genetics, mutations are changes in a genomic sequence: the DNA sequence of a cell's genome or the DNA or RNA sequence of a virus. They can be defined as sudden and spontaneous changes in the cell. Mutations are caused by radiation, viruses, transposons and mutagenic...

s, but by the non-sexual fusion of genetically distinct fungal hyphae. This can lead to individuals that have different nuclei in different parts of their mycelium. In this case, the notion of individual itself becomes vague since the rule of one genome = one individual does not apply any more.

Heterokaryosis is most common in fungi, and lichen
Lichen
Lichens are composite organisms consisting of a symbiotic organism composed of a fungus with a photosynthetic partner , usually either a green alga or cyanobacterium...

, but also occurs in slime molds. This happens because the nuclei in the 'plasmodium' form are the products of many pairwise fusions between amoeboid
Amoeboid
Amoeboids are single-celled life-forms characterized by an irregular shape."Amoeboid" and "amœba" are often used interchangeably even by biologists, and especially refer to a creature moving by using pseudopodia. Most references to "amoebas" or "amoebae" are to amoeboids in general rather than to...

 haploid individuals.
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