Extranuclear inheritance
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Extranuclear inheritance or cytoplasmic inheritance is the transmission of genes that occur outside the
nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: * Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucl ...
. It is found in most
eukaryotes 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. Bacter ...
and is commonly known to occur in cytoplasmic
organelles In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
such as
mitochondria A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...
and
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it i ...
s or from cellular parasites like
virus A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Since Dmitri Ivanovsk ...
es or
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 ...
.


Organelles

Mitochondria are organelles which function to transform energy as a result of
cellular respiration Cellular respiration is the process by which biological fuels are oxidised in the presence of an inorganic electron acceptor such as oxygen to produce large amounts of energy, to drive the bulk production of ATP. Cellular respiration may be des ...
. Chloroplasts are organelles which function to produce sugars via
photosynthesis 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 ...
in plants and algae. The
genes In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
located in
mitochondria A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...
and chloroplasts are very important for proper cellular function. The
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
and other extranuclear types of DNA replicate independently of the DNA located in the nucleus, which is typically arranged in chromosomes that only replicate one time preceding cellular division. The extranuclear genomes of mitochondria and chloroplasts however replicate independently of cell division. They replicate in response to a cell's increasing energy needs which adjust during that cell's lifespan. Since they replicate independently, genomic recombination of these genomes is rarely found in offspring, contrary to nuclear genomes in which recombination is common. Mitochondrial diseases are inherited from the mother, not from the father. Mitochondria with their mitochondrial DNA are already present in the egg cell before it gets fertilized by a sperm. In many cases of fertilization, the head of the sperm enters the egg cell; leaving its middle part, with its mitochondria, behind. The mitochondrial DNA of the sperm often remains outside the
zygote A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism. In multicell ...
and gets excluded from inheritance.


Parasites

Extranuclear transmission of viral genomes and symbiotic bacteria is also possible. An example of viral genome transmission is perinatal transmission. This occurs from mother to fetus during the perinatal period, which begins before birth and ends about 1 month after birth. During this time viral material may be passed from mother to child in the bloodstream or breastmilk. This is of particular concern with mothers carrying HIV or
Hepatitis C Hepatitis C is an infectious disease caused by the hepatitis C virus (HCV) that primarily affects the liver; it is a type of viral hepatitis. During the initial infection people often have mild or no symptoms. Occasionally a fever, dark urine, ...
viruses. Symbiotic cytoplasmic bacteria are also inherited in organisms such as insects and protists.


Types

Three general types of extranuclear inheritance exist. *Vegetative segregation results from random replication and partitioning of cytoplasmic organelles. It occurs with chloroplasts and mitochondria during mitotic cell divisions and results in daughter cells that contain a random sample of the parent cell's organelles. An example of vegetative segregation is with mitochondria of asexually replicating yeast cells. *Uniparental inheritance occurs in extranuclear genes when only one parent contributes organellar DNA to the offspring. A classic example of uniparental gene transmission is the maternal inheritance of human mitochondria. The mother's mitochondria are transmitted to the offspring at
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Pro ...
via the egg. The father's mitochondrial genes are not transmitted to the offspring via the sperm. Very rare cases which require further investigation have been reported of paternal mitochondrial inheritance in humans, in which the father's mitochondrial genome is found in offspring. Chloroplast genes can also inherit uniparentally during
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
. They are historically thought to inherit maternally, but paternal inheritance in many species is increasingly being identified. The mechanisms of uniparental inheritance from species to species differ greatly and are quite complicated. For instance, chloroplasts have been found to exhibit maternal, paternal and biparental modes even within the same species. *Biparental inheritance occurs in extranuclear genes when both parents contribute organellar DNA to the offspring. It may be less common than uniparental extranuclear inheritance, and usually occurs in a permissible species only a fraction of the time. An example of biparental mitochondrial inheritance is in the
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 ...
''
Saccharomyces cerevisiae ''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungus microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have b ...
''. When two
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, respective ...
cells of opposite mating type fuse they can both contribute mitochondria to the resulting
diploid 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, respectiv ...
offspring.


Mutant mitochondria

''Poky'' is a mutant of the fungus ''
Neurospora crassa ''Neurospora crassa'' is a type of red bread mold of the phylum Ascomycota. The genus name, meaning "nerve spore" in Greek, refers to the characteristic striations on the spores. The first published account of this fungus was from an infestation ...
'' that has extranuclear inheritance. ''Poky'' is characterized by slow growth, a defect in mitochondrial ribosome assembly and deficiencies in several
cytochrome Cytochromes are redox-active proteins containing a heme, with a central Fe atom at its core, as a cofactor. They are involved in electron transport chain and redox catalysis. They are classified according to the type of heme and its mode of ...
s. The studies of ''poky'' mutants were among the first to establish an extranuclear mitochondrial basis for inheritance of a particular genotype. It was initially found, using genetic crosses, that ''poky'' is maternally inherited. Subsequently, the primary defect in the ''poky'' mutants was determined to be a deletion in the
mitochondrial A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used t ...
DNA sequence encoding the small subunit of mitochondrial
ribosomal RNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from riboso ...
.


See also

*
Maternal effect A maternal effect is a situation where the phenotype of an organism is determined not only by the environment it experiences and its genotype, but also by the environment and genotype of its mother. In genetics, maternal effects occur when an or ...
*
Plasmid A plasmid is a small, extrachromosomal DNA molecule within a cell that is physically separated from chromosomal DNA and can replicate independently. They are most commonly found as small circular, double-stranded DNA molecules in bacteria; how ...
*
Xenia (plants) Xenia (also known as the Xenia effect) in plants is the effect of pollen on seeds and fruit of the fertilized plant. The effect is separate from the contribution of the pollen towards the next generation. The term was coined in 1881 by the botanis ...


References

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External links

* https://web.archive.org/web/20080521093531/http://www.tamu.edu/classes/magill/gene603/Lecture%20outlines/cytoplasmic%20inh/CYTOPLASMIC_INHERITANCE.html Genetics