Cajal bodies (CBs), also coiled bodies, are spherical

nuclear bodies Nuclear bodies (also known as nuclear domains or nuclear dots) are biomolecular condensates, membraneless structures found in the Cell nucleus, cell nuclei of eukaryote, eukaryotic cell (biology), cells. Nuclear bodies include Cajal body, Cajal b ...

of 0.3–1.0 μm in diameter found in 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 Nucleu ...

of proliferative

cell Cell most often refers to: * Cell (biology), the functional basic unit of life * Cellphone, a phone connected to a cellular network * Clandestine cell, a penetration-resistant form of a secret or outlawed organization * Electrochemical cell, a de ...

s like

embryonic cell In biology, a blastomere is a type of cell produced by cell division (cleavage) of the zygote after fertilization; blastomeres are an essential part of blastula formation, and blastocyst formation in mammals. Human blastomere characteristics In ...

s and tumor cells, or metabolically active cells like

neuron A neuron (American English), neurone (British English), or nerve cell, is an membrane potential#Cell excitability, excitable cell (biology), cell that fires electric signals called action potentials across a neural network (biology), neural net ...

s. CBs are membrane-less

organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as Organ (anatomy), organs are to th ...

s and largely consist of

protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...

s and

RNA Ribonucleic acid (RNA) is a polymeric molecule that is essential for most biological functions, either by performing the function itself (non-coding RNA) or by forming a template for the production of proteins (messenger RNA). RNA and deoxyrib ...

. They were first reported by

Santiago Ramón y Cajal Santiago Ramón y Cajal (; 1 May 1852 – 17 October 1934) was a Spanish neuroscientist, pathologist, and histologist specializing in neuroanatomy, and the central nervous system. He and Camillo Golgi received the Nobel Prize in Physiology or M ...

in 1903, who called them ''nucleolar accessory bodies'' due to their association with the

nucleoli The nucleolus (; : nucleoli ) is the largest structure in the nucleus of eukaryotic cells. It is best known as the site of ribosome biogenesis. The nucleolus also participates in the formation of signal recognition particles and plays a ro ...

in neuronal cells. They were rediscovered with the use of the

electron microscope An electron microscope is a microscope that uses a beam of electrons as a source of illumination. It uses electron optics that are analogous to the glass lenses of an optical light microscope to control the electron beam, for instance focusing it ...

(EM) and named ''coiled bodies'', according to their appearance as coiled threads on EM images, and later renamed after their discoverer. Research on CBs was accelerated after discovery and

cloning Cloning is the process of producing individual organisms with identical genomes, either by natural or artificial means. In nature, some organisms produce clones through asexual reproduction; this reproduction of an organism by itself without ...

of the marker protein p80/Coilin. CBs have been implicated in RNA-related metabolic processes such as the biogenesis, maturation and recycling of

snRNP snRNPs (pronounced "snurps"), or small nuclear ribonucleoproteins, are RNA-protein complexes that combine with unmodified pre-mRNA and various other proteins to form a spliceosome, a large RNA-protein molecular complex upon which splicing of pre- ...

histone In biology, histones are highly basic proteins abundant in lysine and arginine residues that are found in eukaryotic cell nuclei and in most Archaeal phyla. They act as spools around which DNA winds to create structural units called nucleosomes ...

mRNA processing 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 ...

maintenance. CBs assemble RNA which is used by

telomerase Telomerase, also called terminal transferase, is a ribonucleoprotein that adds a species-dependent telomere repeat sequence to the 3' end of telomeres. A telomere is a region of repetitive sequences at each end of the chromosomes of most euka ...

to add

nucleotide Nucleotides are Organic compound, organic molecules composed of a nitrogenous base, a pentose sugar and a phosphate. They serve as monomeric units of the nucleic acid polymers – deoxyribonucleic acid (DNA) and ribonucleic acid (RNA), both o ...

s to the ends of telomeres.

History

CBs were initially discovered by neurobiologist

in 1903 as small argyrophilic (readily stained by silver salts, literally "silver loving") spots in the nuclei of silver-stained neuronal cells. Because of their close association with nucleoli, he named them ''nucleolar accessory bodies''. Later on, they were forgotten and rediscovered multiple times independently which led to a state where scientists from different research fields used different names for the same structure. Names used for CBs included "sphere organelles", "Binnenkörper", "nucleolar bodies" or "coiled bodies". The name coiled bodies comes from observation of electron microscopists Monneron and Bernhard. They described bodies as aggregates composed of coiled threads with thickness of 400–600 Å. When using higher magnification, they appear as tiny, 50 Å thick fibrils irregularly twisted along the axis of the threads. The bodies were even predicted to consist of

ribonucleoproteins Nucleoproteins are proteins conjugated with nucleic acids (either DNA or RNA). Typical nucleoproteins include ribosomes, nucleosomes and viral nucleocapsid proteins. Structures Nucleoproteins tend to be positively charged, facilitating inter ...

since treatment of cells with

protease A protease (also called a peptidase, proteinase, or proteolytic enzyme) is an enzyme that catalysis, catalyzes proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the formation of new protein products ...

and

RNase Ribonuclease (commonly abbreviated RNase) is a type of nuclease that catalyzes the degradation of RNA into smaller components. Ribonucleases can be divided into endoribonucleases and exoribonucleases, and comprise several sub-classes within the ...

together, but not alone, caused dramatic changes to the structure of CBs.

Localization

Cajal bodies are found in all eukaryotes that have been carefully studied. The cells in which Cajal bodies are most apparent usually demonstrate high levels of transcriptional activity, and are often dividing rapidly.

Cell cycle

They are about 0.1–2.0

micrometre The micrometre (English in the Commonwealth of Nations, Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a uni ...

s and are found one to five per nucleus. The number varies in different types of cells and over the cell cycle. Maximum number is reached in mid G₁ phase and towards G₂ they become larger and their number decreases. CBs disassemble during the M phase and reappear again later in G₁ phase. Cajal bodies are possibly sites of assembly or modification of the transcription machinery of the nucleus.

Functions

CBs are bound to 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 ...

by coilin proteins. P80-coilin is a specific marker for coiled bodies, and demonstrates these bodies tend to be associated with the nucleolus when cells are not dividing. CBs are associated with telomerase assembly and recruitment via a CAB-RNA sequence common in both CB RNAs ( scaRNAs) and the RNA component of telomerase (TERC). TCAB1 recognizes the CAB sequence in both and recruits telomerase to the CBs. CBs contain high concentrations of splicing small nuclear ribonucleoproteins (

s), possibly indicating that they function to modify RNA after it has been transcribed from DNA. Experimental evidence indicates that CBs contribute to the biogenesis of enzyme

, and assist in subsequent transport of telomerase to

References

{{Nucleus Organelles S Cell nucleus Cell anatomy Telomeres