] Nucleolus organizer regions (NORs) are chromosome, chromosomal regions crucial for the formation of the

nucleolus The nucleolus (; : nucleoli ) is the largest structure in the cell nucleus, nucleus of eukaryote, eukaryotic cell (biology), cells. It is best known as the site of ribosome biogenesis. The nucleolus also participates in the formation of signa ...

. In humans, the NORs are located on the short arms of the acrocentric chromosomes 13, 14, 15, 21 and 22, the genes RNR1, RNR2, RNR3, RNR4, and RNR5 respectively. These regions code for 5.8S, 18S, and 28S

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 ribosomal ...

. The NORs are "sandwiched" between the repetitive, heterochromatic

DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...

sequences of the

centromere The centromere links a pair of sister chromatids together during cell division. This constricted region of chromosome connects the sister chromatids, creating a short arm (p) and a long arm (q) on the chromatids. During mitosis, spindle fiber ...

s and

telomere A telomere (; ) is a region of repetitive nucleotide sequences associated with specialized proteins at the ends of linear chromosomes (see #Sequences, Sequences). Telomeres are a widespread genetic feature most commonly found in eukaryotes. In ...

s. The exact sequence of these regions is not included in the human reference genome as of 2016 or the GRCh38.p10 released January 6, 2017. On 28 February 2019, GRCh38.p13 was released, which added the NOR sequences for the short arms of chromosomes 13, 14, 15, 21, and 22. However, it is known that NORs contain tandem copies of

ribosomal DNA The ribosomal DNA (rDNA) consists of a group of ribosomal RNA encoding genes and related regulatory elements, and is widespread in similar configuration in all domains of life. The ribosomal DNA encodes the non-coding ribosomal RNA, integral struc ...

(rDNA) genes. Some sequences of flanking sequences proximal and distal to NORs have been reported. The NORs of a

loris Loris is the common name for the strepsirrhine mammals of the subfamily Lorinae (sometimes spelled Lorisinae) in the family Lorisidae. ''Loris'' is one genus in this subfamily and includes the slender lorises, ''Nycticebus'' is the genus cont ...

have been reported to be highly variable. There are also DNA sequences related to rDNA that are on other chromosomes and may be involved in nucleoli formation. NORAgc

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]

Barbara McClintock Barbara McClintock (June 16, 1902 – September 2, 1992) was an American scientist and cytogenetics, cytogeneticist who was awarded the 1983 Nobel Prize in Physiology or Medicine. McClintock received her PhD in botany from Cornell University ...

first described the "nucleolar-organizing body" in ''

Zea mays Maize (; ''Zea mays''), also known as corn in North American English, is a tall stout Poaceae, grass that produces cereal grain. It was domesticated by indigenous peoples of Mexico, indigenous peoples in southern Mexico about 9,000 years ago ...

'' in 1934. In

karyotype A karyotype is the general appearance of the complete set of chromosomes in the cells of a species or in an individual organism, mainly including their sizes, numbers, and shapes. Karyotyping is the process by which a karyotype is discerned by de ...

analysis, a silver stain can be used to identify the NOR. NORs can also be seen in nucleoli using silver stain, and that is being used to investigate cancerous changes. NORs can also be seen using

antibodies An antibody (Ab) or immunoglobulin (Ig) is a large, Y-shaped protein belonging to the immunoglobulin superfamily which is used by the immune system to identify and neutralize antigens such as bacteria and viruses, including those that caus ...

directed against the protein UBF, which binds to NOR DNA.

Molecular biology

In addition to UBF, NORs also bind to ATRX protein,

treacle Treacle () is any uncrystallised syrup made during the refining of sugar.Oxford Dictionary The most common forms of treacle are golden syrup, a pale variety, and black treacle, a darker variety similar to molasses. Black treacle has a distinctiv ...

, sirtuin-7 and other proteins. UBF has been identified as a mitotic "bookmark" of expressed rDNA, which allows it to resume transcription quickly after

mitosis Mitosis () is a part of the cell cycle in eukaryote, eukaryotic cells in which replicated chromosomes are separated into two new Cell nucleus, nuclei. Cell division by mitosis is an equational division which gives rise to genetically identic ...

. The distal flanking junction (DJ) of the NORs has been shown to associate with the periphery of nucleoli. rDNA operons in ''

Escherichia coli ''Escherichia coli'' ( )Wells, J. C. (2000) Longman Pronunciation Dictionary. Harlow ngland Pearson Education Ltd. is a gram-negative, facultative anaerobic, rod-shaped, coliform bacterium of the genus '' Escherichia'' that is commonly fo ...

'' have been found to cluster near each other, similar to a eukaryotic nucleolus.

References

{{DEFAULTSORT:Nucleolus Organizer Region Cell anatomy Molecular genetics Nuclear substructures

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Molecular biology

See also

References