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

nucleolus The nucleolus (, plural: nucleoli ) is the largest structure in the nucleus of eukaryotic cells. It is best known as the site of ribosome biogenesis, which is the synthesis of ribosomes. The nucleolus also participates in the formation of s ...

. In humans, the NORs are located on the short arms of the

acrocentric 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 fibers ...

chromosomes 13, 14, 15, 21 and 22, the genes RNR1, RNR2, RNR3, RNR4, and RNR5 respectively. These regions code for

5.8S In molecular biology, the 5.8S ribosomal RNA (5.8S rRNA) is a non-coding RNA component of the large subunit of the eukaryotic ribosome and so plays an important role in protein translation. It is transcribed by RNA polymerase I as part of the 4 ...

, 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 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 fibers ...

s and

telomere A telomere (; ) is a region of repetitive nucleotide sequences associated with specialized proteins at the ends of linear chromosomes. Although there are different architectures, telomeres, in a broad sense, are a widespread genetic feature mo ...

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 Ribosomal DNA (rDNA) is a DNA sequence that codes for ribosomal RNA. These sequences regulate transcription initiation and amplification, and contain both transcribed and non-transcribed spacer segments. In the human genome there are 5 chrom ...

(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 con ...

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 cytogeneticist who was awarded the 1983 Nobel Prize in Physiology or Medicine. McClintock received her PhD in botany from Cornell University in 1927. There ...

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

Zea mays Maize ( ; ''Zea mays'' subsp. ''mays'', from es, maíz after tnq, mahiz), also known as corn ( North American and Australian English), is a cereal grain first domesticated by indigenous peoples in southern Mexico about 10,000 years ago. ...

'' in 1934. In

karyotype A karyotype is the general appearance of the complete set of metaphase 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 disce ...

analysis, a

silver stain In pathology, silver staining is the use of silver to selectively alter the appearance of a target in microscopy of histological sections; in temperature gradient gel electrophoresis; and in polyacrylamide gels. In traditional stained glass, si ...

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 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 a darker variety known as black treacle, similar to molasses. Black treacle has ...

, 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 In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is maint ...

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

operon In genetics, an operon is a functioning unit of DNA containing a cluster of genes under the control of a single promoter. The genes are transcribed together into an mRNA strand and either translated together in the cytoplasm, or undergo spli ...

s in ''

Escherichia coli ''Escherichia coli'' (),Wells, J. C. (2000) Longman Pronunciation Dictionary. Harlow ngland Pearson Education Ltd. also known as ''E. coli'' (), is a Gram-negative, facultative anaerobic, rod-shaped, coliform bacterium of the genus '' Esc ...

'' 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