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Because the nuclear membrane is impermeable to large molecules , nuclear pores are required that regulate nuclear transport of molecules across the envelope . The pores cross both nuclear membranes , providing a channel through which larger molecules must be actively transported by carrier proteins while allowing free...
= = History = =
The nucleus was the first organelle to be discovered . What is most likely the oldest preserved drawing dates back to the early microscopist Antonie van Leeuwenhoek ( 1632 – 1723 ) . He observed a " lumen " , the nucleus , in the red blood cells of salmon . Unlike mammalian red blood cells , those of other vertebrates...
The nucleus was also described by Franz Bauer in 1804 and in more detail in 1831 by Scottish botanist Robert Brown in a talk at the Linnean Society of London . Brown was studying orchids under microscope when he observed an opaque area , which he called the " areola " or " nucleus " , in the cells of the flower 's out...
He did not suggest a potential function . In 1838 , Matthias Schleiden proposed that the nucleus plays a role in generating cells , thus he introduced the name " cytoblast " ( cell builder ) . He believed that he had observed new cells assembling around " cytoblasts " . Franz Meyen was a strong opponent of this view ,...
Between 1877 and 1878 , Oscar Hertwig published several studies on the fertilization of sea urchin eggs , showing that the nucleus of the sperm enters the oocyte and fuses with its nucleus . This was the first time it was suggested that an individual develops from a ( single ) nucleated cell . This was in contradictio...
= = Structures = =
The nucleus is the largest cellular organelle in animal cells . In mammalian cells , the average diameter of the nucleus is approximately 6 micrometres ( µm ) , which occupies about 10 % of the total cell volume . The viscous liquid within it is called nucleoplasm , and is similar in composition to the cytosol found o...
= = = Nuclear envelope and pores = = =
The nuclear envelope , otherwise known as nuclear membrane , consists of two cellular membranes , an inner and an outer membrane , arranged parallel to one another and separated by 10 to 50 nanometres ( nm ) . The nuclear envelope completely encloses the nucleus and separates the cell 's genetic material from the surr...
Nuclear pores , which provide aqueous channels through the envelope , are composed of multiple proteins , collectively referred to as nucleoporins . The pores are about 125 million daltons in molecular weight and consist of around 50 ( in yeast ) to several hundred proteins ( in vertebrates ) . The pores are 100 nm in...
Most proteins , ribosomal subunits , and some DNAs are transported through the pore complexes in a process mediated by a family of transport factors known as karyopherins . Those karyopherins that mediate movement into the nucleus are also called importins , whereas those that mediate movement out of the nucleus are c...
= = = Nuclear lamina = = =
In animal cells , two networks of intermediate filaments provide the nucleus with mechanical support : The nuclear lamina forms an organized meshwork on the internal face of the envelope , while less organized support is provided on the cytosolic face of the envelope . Both systems provide structural support for the n...
The nuclear lamina is composed mostly of lamin proteins . Like all proteins , lamins are synthesized in the cytoplasm and later transported to the nucleus interior , where they are assembled before being incorporated into the existing network of nuclear lamina . Lamins found on the cytosolic face of the membrane , suc...
Like the components of other intermediate filaments , the lamin monomer contains an alpha @-@ helical domain used by two monomers to coil around each other , forming a dimer structure called a coiled coil . Two of these dimer structures then join side by side , in an antiparallel arrangement , to form a tetramer calle...
Mutations in lamin genes leading to defects in filament assembly cause a group of rare genetic disorders known as laminopathies . The most notable laminopathy is the family of diseases known as progeria , which causes the appearance of premature aging in its sufferers . The exact mechanism by which the associated bioc...
= = = Chromosomes = = =
The cell nucleus contains the majority of the cell 's genetic material in the form of multiple linear DNA molecules organized into structures called chromosomes . Each human cell contains roughly two meters of DNA . During most of the cell cycle these are organized in a DNA @-@ protein complex known as chromatin , and...
There are two types of chromatin . Euchromatin is the less compact DNA form , and contains genes that are frequently expressed by the cell . The other type , heterochromatin , is the more compact form , and contains DNA that is infrequently transcribed . This structure is further categorized into facultative heterochr...
Antibodies to certain types of chromatin organization , in particular , nucleosomes , have been associated with a number of autoimmune diseases , such as systemic lupus erythematosus . These are known as anti @-@ nuclear antibodies ( ANA ) and have also been observed in concert with multiple sclerosis as part of gener...
= = = Nucleolus = = =
The nucleolus is a discrete densely stained structure found in the nucleus . It is not surrounded by a membrane , and is sometimes called a suborganelle . It forms around tandem repeats of rDNA , DNA coding for ribosomal RNA ( rRNA ) . These regions are called nucleolar organizer regions ( NOR ) . The main roles of th...
In the first step of ribosome assembly , a protein called RNA polymerase I transcribes rDNA , which forms a large pre @-@ rRNA precursor . This is cleaved into the subunits 5.8S , 18S , and 28S rRNA . The transcription , post @-@ transcriptional processing , and assembly of rRNA occurs in the nucleolus , aided by smal...
When observed under the electron microscope , the nucleolus can be seen to consist of three distinguishable regions : the innermost fibrillar centers ( FCs ) , surrounded by the dense fibrillar component ( DFC ) , which in turn is bordered by the granular component ( GC ) . Transcription of the rDNA occurs either in t...
= = = Other subnuclear bodies = = =
Besides the nucleolus , the nucleus contains a number of other non @-@ membrane @-@ delineated bodies . These include Cajal bodies , Gemini of coiled bodies , polymorphic interphase karyosomal association ( PIKA ) , promyelocytic leukaemia ( PML ) bodies , paraspeckles , and splicing speckles . Although little is know...
Other subnuclear structures appear as part of abnormal disease processes . For example , the presence of small intranuclear rods has been reported in some cases of nemaline myopathy . This condition typically results from mutations in actin , and the rods themselves consist of mutant actin as well as other cytoskeleta...
= = = = Cajal bodies and gems = = = =
A nucleus typically contains between 1 and 10 compact structures called Cajal bodies or coiled bodies ( CB ) , whose diameter measures between 0 @.@ 2 µm and 2 @.@ 0 µm depending on the cell type and species . When seen under an electron microscope , they resemble balls of tangled thread and are dense foci of distribu...
Similar to Cajal bodies are Gemini of Cajal bodies , or gems , whose name is derived from the Gemini constellation in reference to their close " twin " relationship with CBs . Gems are similar in size and shape to CBs , and in fact are virtually indistinguishable under the microscope . Unlike CBs , gems do not contain...
= = = = RAFA and PTF domains = = = =
RAFA domains , or polymorphic interphase karyosomal associations , were first described in microscopy studies in 1991 . Their function remains unclear , though they were not thought to be associated with active DNA replication , transcription , or RNA processing . They have been found to often associate with discrete ...
= = = = PML bodies = = = =
Promyelocytic leukaemia bodies ( PML bodies ) are spherical bodies found scattered throughout the nucleoplasm , measuring around 0 @.@ 1 – 1 @.@ 0 µm . They are known by a number of other names , including nuclear domain 10 ( ND10 ) , Kremer bodies , and PML oncogenic domains . PML bodies are named after one of their ...
= = = = Splicing speckles = = = =
Speckles are subnuclear structures that are enriched in pre @-@ messenger RNA splicing factors and are located in the interchromatin regions of the nucleoplasm of mammalian cells . At the fluorescence @-@ microscope level they appear as irregular , punctate structures , which vary in size and shape , and when examined...
= = = = Paraspeckles = = = =
Discovered by Fox et al. in 2002 , paraspeckles are irregularly shaped compartments in the nucleus ' interchromatin space . First documented in HeLa cells , where there are generally 10 – 30 per nucleus , paraspeckles are now known to also exist in all human primary cells , transformed cell lines , and tissue sections...
Paraspeckles are dynamic structures that are altered in response to changes in cellular metabolic activity . They are transcription dependent and in the absence of RNA Pol II transcription , the paraspeckle disappears and all of its associated protein components ( PSP1 , p54nrb , PSP2 , CFI ( m ) 68 , and PSF ) form a...
= = = = Perichromatin fibrils = = = =
Perichromatin fibrils are visible only under electron microscope . They are located next to the transcriptionally active chromatin and are hypothesized to be the sites of active pre @-@ mRNA processing .
= = Function = =
The nucleus provides a site for genetic transcription that is segregated from the location of translation in the cytoplasm , allowing levels of gene regulation that are not available to prokaryotes . The main function of the cell nucleus is to control gene expression and mediate the replication of DNA during the cell ...
The nucleus is an organelle found in eukaryotic cells . Inside its fully enclosed nuclear membrane , it contains the majority of the cell 's genetic material . This material is organized as DNA molecules , along with a variety of proteins , to form chromosomes .
= = = Cell compartmentalization = = =
The nuclear envelope allows the nucleus to control its contents , and separate them from the rest of the cytoplasm where necessary . This is important for controlling processes on either side of the nuclear membrane . In most cases where a cytoplasmic process needs to be restricted , a key participant is removed to th...
In order to control which genes are being transcribed , the cell separates some transcription factor proteins responsible for regulating gene expression from physical access to the DNA until they are activated by other signaling pathways . This prevents even low levels of inappropriate gene expression . For example , ...
The compartmentalization allows the cell to prevent translation of unspliced mRNA . Eukaryotic mRNA contains introns that must be removed before being translated to produce functional proteins . The splicing is done inside the nucleus before the mRNA can be accessed by ribosomes for translation . Without the nucleus ,...
= = = Gene expression = = =
Gene expression first involves transcription , in which DNA is used as a template to produce RNA . In the case of genes encoding proteins , that RNA produced from this process is messenger RNA ( mRNA ) , which then needs to be translated by ribosomes to form a protein . As ribosomes are located outside the nucleus , m...
Since the nucleus is the site of transcription , it also contains a variety of proteins that either directly mediate transcription or are involved in regulating the process . These proteins include helicases , which unwind the double @-@ stranded DNA molecule to facilitate access to it , RNA polymerases , which synthe...
= = = Processing of pre @-@ mRNA = = =
Newly synthesized mRNA molecules are known as primary transcripts or pre @-@ mRNA . They must undergo post @-@ transcriptional modification in the nucleus before being exported to the cytoplasm ; mRNA that appears in the cytoplasm without these modifications is degraded rather than used for protein translation . The t...
RNA splicing , carried out by a complex called the spliceosome , is the process by which introns , or regions of DNA that do not code for protein , are removed from the pre @-@ mRNA and the remaining exons connected to re @-@ form a single continuous molecule . This process normally occurs after 5 ' capping and 3 ' po...
= = Dynamics and regulation = =
= = = Nuclear transport = = =
The entry and exit of large molecules from the nucleus is tightly controlled by the nuclear pore complexes . Although small molecules can enter the nucleus without regulation , macromolecules such as RNA and proteins require association karyopherins called importins to enter the nucleus and exportins to exit . " Cargo...
Nuclear import depends on the importin binding its cargo in the cytoplasm and carrying it through the nuclear pore into the nucleus . Inside the nucleus , RanGTP acts to separate the cargo from the importin , allowing the importin to exit the nucleus and be reused . Nuclear export is similar , as the exportin binds th...
Specialized export proteins exist for translocation of mature mRNA and tRNA to the cytoplasm after post @-@ transcriptional modification is complete . This quality @-@ control mechanism is important due to these molecules ' central role in protein translation . Mis @-@ expression of a protein due to incomplete excisio...
= = = Assembly and disassembly = = =