1180:(PML-nuclear 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-nuclear bodies are named after one of their major components, the promyelocytic leukemia protein (PML). They are often seen in the nucleus in association with Cajal bodies and cleavage bodies. Pml-/- mice, which are unable to create PML-nuclear bodies, develop normally without obvious ill effects, showing that PML-nuclear bodies are not required for most essential biological processes.
588:. Together, these membranes serve to separate the cell's genetic material from the rest of the cell contents, and allow the nucleus to maintain an environment distinct from the rest of the cell. Despite their close apposition around much of the nucleus, the two membranes differ substantially in shape and contents. The inner membrane surrounds the nuclear content, providing its defining edge. Embedded within the inner membrane, various proteins bind the intermediate filaments that give the nucleus its structure. The outer membrane encloses the inner membrane, and is continuous with the adjacent
2047:
471:
33:
1034:(SMN) whose function relates to snRNP biogenesis. Gems are believed to assist CBs in snRNP biogenesis, though it has also been suggested from microscopy evidence that CBs and gems are different manifestations of the same structure. Later ultrastructural studies have shown gems to be twins of Cajal bodies with the difference being in the coilin component; Cajal bodies are SMN positive and coilin positive, and gems are SMN positive and coilin negative.
2059:
995:
553:
1401:
85:
1516:
872:. The structural cohesion of the nucleolus depends on its activity, as ribosomal assembly in the nucleolus results in the transient association of nucleolar components, facilitating further ribosomal assembly, and hence further association. This model is supported by observations that inactivation of rDNA results in intermingling of nucleolar structures.
351:
1669:. The sister chromatids can then be pulled to separate locations in the cell. In many cells, the centrosome is located in the cytoplasm, outside the nucleus; the microtubules would be unable to attach to the chromatids in the presence of the nuclear envelope. Therefore, the early stages in the cell cycle, beginning in prophase and until around
2038:, suggests that the nucleus instead originated from a single ancestral cell that evolved a second exterior cell membrane; the interior membrane enclosing the original cell then became the nuclear membrane and evolved increasingly elaborate pore structures for passage of internally synthesized cellular components such as ribosomal subunits.
945:). Transcription of the rDNA occurs either in the FC or at the FC-DFC boundary, and, therefore, when rDNA transcription in the cell is increased, more FCs are detected. Most of the cleavage and modification of rRNAs occurs in the DFC, while the latter steps involving protein assembly onto the ribosomal subunits occur in the GC.
643:). The pores are 100 nm in total diameter; however, the gap through which molecules freely diffuse is only about 9 nm wide, due to the presence of regulatory systems within the center of the pore. This size selectively allows the passage of small water-soluble molecules while preventing larger molecules, such as
2007:, posits that the membrane-bound nucleus, along with other eukaryotic features, originated from the infection of a prokaryote by a virus. The suggestion is based on similarities between eukaryotes and viruses such as linear DNA strands, mRNA capping, and tight binding to proteins (analogizing histones to
1336:
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.
1312:
allows control of the nuclear contents, and separates 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 the nucleus,
1194:
Discovered by Fox et al. in 2002, paraspeckles are irregularly shaped compartments in the interchromatin space of the nucleus. 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
1413:
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
1708:
for early apoptotic activity. Cells that express mutant caspase-resistant lamins are deficient in nuclear changes related to apoptosis, suggesting that lamins play a role in initiating the events that lead to apoptotic degradation of the nucleus. Inhibition of lamin assembly itself is an inducer of
1695:
Apoptosis is a controlled process in which the cell's structural components are destroyed, resulting in death of the cell. Changes associated with apoptosis directly affect the nucleus and its contents, for example, in the condensation of chromatin and the disintegration of the nuclear envelope and
1656:
At a certain point during the cell cycle in open mitosis, the cell divides to form two cells. In order for this process to be possible, each of the new daughter cells must have a full set of genes, a process requiring replication of the chromosomes as well as segregation of the separate sets. This
953:
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
1487:
connected to re-form a single continuous molecule. This process normally occurs after 5' capping and 3' polyadenylation but can begin before synthesis is complete in transcripts with many exons. Many pre-mRNAs can be spliced in multiple ways to produce different mature mRNAs that encode different
1375:
The main function of the cell nucleus is to control gene expression and mediate the replication of DNA during the cell cycle. It has been found that replication happens in a localised way in the cell nucleus. In the S phase of interphase of the cell cycle; replication takes place. Contrary to the
1164:
PIKA 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
985:
embryos. B snurposomes appear alone or attached to the Cajal bodies in the electron micrographs of the amphibian nuclei. While nuclear speckles were originally thought to be storage sites for the splicing factors, a more recent study demonstrated that organizing genes and pre-mRNA substrates near
1684:, the nuclear envelope remains intact, the centrosomes are located in the cytoplasm, and the microtubules come in contact with chromosomes, whose centromeric regions are incorporated into the nuclear envelope (the so-called closed mitosis with extranuclear spindle). In many other protists (e.g.,
1582:
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
1578:
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
1198:
Paraspeckles sequester nuclear proteins and RNA and thus appear to function as a molecular sponge that is involved in the regulation of gene expression. Furthermore, paraspeckles are dynamic structures that are altered in response to changes in cellular metabolic activity. They are transcription
1360:
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.
1199:
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 crescent shaped perinucleolar cap in the nucleolus. This phenomenon is demonstrated during the cell cycle. In the
724:
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, such as
716:
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 nuclear envelope and anchoring sites for chromosomes and nuclear pores.
1592:
1575:, which is bound to either GTP or GDP (guanosine diphosphate), depending on whether it is located in the nucleus or the cytoplasm. Whereas importins depend on RanGTP to dissociate from their cargo, exportins require RanGTP in order to bind to their cargo.
1408:
during transcription, highlighting the possibility of transcribing more than one gene at a time. The diagram includes 8 RNA polymerases however the number can vary depending on cell type. The image also includes transcription factors and a porous, protein
1546:
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
1006:
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 distribution for the protein
2125:
was a strong opponent of this view, having already described cells multiplying by division and believing that many cells would have no nuclei. The idea that cells can be generated de novo, by the "cytoblast" or otherwise, contradicted work by
2083:
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
1333:, a molecule made later from glucose-6-phosphate, a regulator protein removes hexokinase to the nucleus, where it forms a transcriptional repressor complex with nuclear proteins to reduce the expression of genes involved in glycolysis.
658:. Those karyopherins that mediate movement into the nucleus are also called importins, whereas those that mediate movement out of the nucleus are called exportins. Most karyopherins interact directly with their cargo, although some use
6499:
The Works of, or arcana of nature by means of exactissimorum microscopes had been detected and confirmed by a variety of experiments, the
Epistles to the various illustrious men of valor J. Arnold and Delphi, A. Beman, Lugdina York
1959:, which are thought to have developed from a similar endosymbiotic relationship between proto-eukaryotes and aerobic bacteria. One possibility is that the nuclear membrane arose as a new membrane system following the origin of
1712:
The nuclear envelope acts as a barrier that prevents both DNA and RNA viruses from entering the nucleus. Some viruses require access to proteins inside the nucleus in order to replicate and/or assemble. DNA viruses, such as
1267:
has also been shown to cause the formation of clastosomes. These nuclear bodies contain catalytic and regulatory subunits of the proteasome and its substrates, indicating that clastosomes are sites for degrading proteins.
1140:
above (e.g., nucleolus, nuclear speckles, Cajal bodies) the nucleus contains a number of other nuclear bodies. These include polymorphic interphase karyosomal association (PIKA), promyelocytic leukaemia (PML) bodies, and
1801:
may induce the release of some immature "micronucleated" erythrocytes into the bloodstream. Anucleated cells can also arise from flawed cell division in which one daughter lacks a nucleus and the other has two nuclei.
1649:, in which the nuclear envelope remains intact. In closed mitosis, the daughter chromosomes migrate to opposite poles of the nucleus, which then divides in two. The cells of higher eukaryotes, however, usually undergo
965:
Studies on the composition, structure and behaviour of speckles have provided a model for understanding the functional compartmentalization of the nucleus and the organization of the gene-expression machinery splicing
607:(NPCs) perforating the envelope. Each NPC contains an eightfold-symmetric ring-shaped structure at a position where the inner and outer membranes fuse. The number of NPCs can vary considerably across cell types; small
958:. Speckles are dynamic structures, and both their protein and RNA-protein components can cycle continuously between speckles and other nuclear locations, including active transcription sites. Speckles can work with
1641:). During these events, the structural components of the nucleus — the envelope and lamina — can be systematically degraded. In most cells, the disassembly of the nuclear envelope marks the end of the
970:
and other splicing proteins necessary for pre-mRNA processing. Because of a cell's changing requirements, the composition and location of these bodies changes according to mRNA transcription and regulation via
1946:
created the nucleus-containing eukaryotic cell. (Organisms of the
Archaeal and Bacterial domains have no cell nucleus.) It is hypothesized that the symbiosis originated when ancient archaea similar to modern
1195:
tissue sections. Their name is derived from their distribution in the nucleus; the "para" is short for parallel and the "speckles" refers to the splicing speckles to which they are always in close proximity.
1583:
excision of exons or mis-incorporation of amino acids could have negative consequences for the cell; thus, incompletely modified RNA that reaches the cytoplasm is degraded rather than used in translation.
1455:
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 three main modifications are
1717:
replicate and assemble in the cell nucleus, and exit by budding through the inner nuclear membrane. This process is accompanied by disassembly of the lamina on the nuclear face of the inner membrane.
1145:. Although little is known about a number of these domains, they are significant in that they show that the nucleoplasm is not a uniform mixture, but rather contains organized functional subdomains.
1026:
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
647:
and larger proteins, from inappropriately entering or exiting the nucleus. These large molecules must be actively transported into the nucleus instead. Attached to the ring is a structure called the
1673:, the nuclear membrane is dismantled. Likewise, during the same period, the nuclear lamina is also disassembled, a process regulated by phosphorylation of the lamins by protein kinases such as the
1417:
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
6495:
Opera Omnia, seu Arcana
Naturae ope exactissimorum Microscopiorum detecta, experimentis variis comprobata, Epistolis ad varios illustres viros J. Arnold et Delphis, A. Beman, Lugdinum Batavorum
4278:"Regional and temporal specialization in the nucleus: a transcriptionally-active nuclear domain rich in PTF, Oct1 and PIKA antigens associates with specific chromosomes early in the cell cycle"
318:
through the pores is required for both gene expression and the maintenance of chromosomes. Although the interior of the nucleus does not contain any membrane-bound subcompartments, a number of
7880:
975:
of specific proteins. The splicing speckles are also known as nuclear speckles (nuclear specks), splicing factor compartments (SF compartments), interchromatin granule clusters (IGCs), and
784:. These filaments can be assembled or disassembled in a dynamic manner, meaning that changes in the length of the filament depend on the competing rates of filament addition and removal.
1645:
of mitosis. However, this disassembly of the nucleus is not a universal feature of mitosis and does not occur in all cells. Some unicellular eukaryotes (e.g., yeasts) undergo so-called
1653:, which is characterized by breakdown of the nuclear envelope. The daughter chromosomes then migrate to opposite poles of the mitotic spindle, and new nuclei reassemble around them.
1931:
origin has been the subject of much speculation. Four major hypotheses have been proposed to explain the existence of the nucleus, although none have yet earned widespread support.
1243:
Clastosomes are small nuclear bodies (0.2–0.5 μm) described as having a thick ring-shape due to the peripheral capsule around these bodies. This name is derived from the Greek
2175:"). Therefore, the necessity of the sperm nucleus for fertilization was discussed for quite some time. However, Hertwig confirmed his observation in other animal groups, including
2110:
under the microscope when he observed an opaque area, which he called the "areola" or "nucleus", in the cells of the flower's outer layer. He did not suggest a potential function.
2171:
of a species would be repeated during embryonic development, including generation of the first nucleated cell from a "monerula", a structureless mass of primordial protoplasm ("
1231:
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
651:
that extends into the nucleoplasm, and a series of filamentous extensions that reach into the cytoplasm. Both structures serve to mediate binding to nuclear transport proteins.
1971:
produced by the protomitochondria. The archaeal origin of the nucleus is supported by observations that archaea and eukarya have similar genes for certain proteins, including
962:
as enhancers of gene activity to directly enhance the activity of certain genes. Moreover, speckle-associating and non-associating p53 gene targets are functionally distinct.
6108:
Bernstein, H., Bernstein, C. (2017). Sexual
Communication in Archaea, the Precursor to Eukaryotic Meiosis. In: Witzany, G. (eds) Biocommunication of Archaea. Springer, Cham.
1915:
are known to have two nuclei. Unlike other multinucleated cells these nuclei contain two distinct lineages of DNA: one from the dinoflagellate and the other from a symbiotic
1769:
An anucleated cell contains no nucleus and is, therefore, incapable of dividing to produce daughter cells. The best-known anucleated cell is the mammalian red blood cell, or
596:
that are actively translating proteins across membrane. The space between the two membranes is called the perinuclear space, and is continuous with the endoplasmic reticulum
1990:
bacteria that possess a nuclear structure with primitive pores and other compartmentalized membrane structures. A similar proposal states that a eukaryote-like cell, the
1704:, which cleave the lamin proteins and, thus, degrade the nucleus' structural integrity. Lamin cleavage is sometimes used as a laboratory indicator of caspase activity in
2542:"The relation of immunoglobulin class, pattern of anti-nuclear antibody, and complement-fixing antibodies to DNA in sera from patients with systemic lupus erythematosus"
1749:
cell types usually have a single nucleus, but some have no nuclei, while others have several. This can result from normal development, as in the maturation of mammalian
5403:
Torous DK, Dertinger SD, Hall NE, Tometsko CR (February 2000). "Enumeration of micronucleated reticulocytes in rat peripheral blood: a flow cytometric study". Primary.
1677:. Towards the end of the cell cycle, the nuclear membrane is reformed, and around the same time, the nuclear lamina are reassembled by dephosphorylating the lamins.
1380:
emerged, which means replication forks are concentrated towards some immobilised 'factory' regions through which the template DNA strands pass like conveyor belts.
526:
is reported across a range of cell types and species. In eukaryotes the nucleus in many cells typically occupies 10% of the cell volume. The nucleus is the largest
2823:
2099:
1148:
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
6846:
A university level textbook focusing on cell biology. Contains information on nucleus structure and function, including nuclear transport, and subnuclear domains
7015:
1313:
where it interacts with transcription factors to downregulate the production of certain enzymes in the pathway. This regulatory mechanism occurs in the case of
1579:
exportin binds the cargo inside the nucleus in a process facilitated by RanGTP, exits through the nuclear pore, and separates from its cargo in the cytoplasm.
733:, bind to the cytoskeleton to provide structural support. Lamins are also found inside the nucleoplasm where they form another regular structure, known as the
2163:
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 contradiction to
654:
Most proteins, ribosomal subunits, and some RNAs are transported through the pore complexes in a process mediated by a family of transport factors known as
1357:
on the NF-κB protein allows it to be transported through the nuclear pore and into the nucleus, where it stimulates the transcription of the target genes.
6776:
A review article about the nucleus, explaining the structure of chromosomes within the organelle, and describing the nucleolus and other subnuclear bodies
4325:
Zimber A, Nguyen QD, Gespach C (October 2004). "Nuclear bodies and compartments: functional roles and cellular signalling in health and disease". Review.
1986:
A second model proposes that proto-eukaryotic cells evolved from bacteria without an endosymbiotic stage. This model is based on the existence of modern
5816:"Tertiary endosymbiosis in two dinotoms has generated little change in the mitochondrial genomes of their dinoflagellate hosts and diatom endosymbionts"
1880:, become multinucleated during development; the resulting arrangement of nuclei near the periphery of the cells allows maximal intracellular space for
433:. Active genes, which are generally found in the euchromatic region of the chromosome, tend to be located towards the chromosome's territory boundary.
1353:, binds to a cell membrane receptor, resulting in the recruitment of signalling proteins, and eventually activating the transcription factor NF-κB. A
830:
1469:
370:
The cell nucleus contains the majority of the cell's genetic material in the form of multiple linear DNA molecules organized into structures called
4465:
2342:
Grigoryev SA, Bulynko YA, Popova EY (2006). "The end adjusts the means: heterochromatin remodelling during terminal cell differentiation". Review.
4822:
Moreno F, Ahuatzi D, Riera A, Palomino CA, Herrero P (February 2005). "Glucose sensing through the Hxk2-dependent signalling pathway". Primary.
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in such a way to promote cell function. The nucleus maintains the integrity of genes and controls the activities of the cell by regulating
2195:
for heredity. The function of the nucleus as carrier of genetic information became clear only later, after mitosis was discovered and the
5296:
Böhm I (November 2007). "IgG deposits can be detected in cell nuclei of patients with both lupus erythematosus and malignancy". Primary.
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Without the nucleus, ribosomes would translate newly transcribed (unprocessed) mRNA, resulting in malformed and nonfunctional proteins.
3936:
6407:
Bell PJ (November 2006). "Sex and the eukaryotic cell cycle is consistent with a viral ancestry for the eukaryotic nucleus". Primary.
1955:, eventually forming the early nucleus. This theory is analogous to the accepted theory for the origin of eukaryotic mitochondria and
1762:
659:
1472:(hnRNPs). Addition of the 5' cap occurs co-transcriptionally and is the first step in post-transcriptional modification. The 3' poly-
7444:
7008:
3066:
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1442:
363:
1483:, 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
2387:"Specific staining of human chromosomes in Chinese hamster x man hybrid cell lines demonstrates interphase chromosome territories"
1555:
to exit. "Cargo" proteins that must be translocated from the cytoplasm to the nucleus contain short amino acid sequences known as
2488:
1789:, where they lose their nuclei, organelles, and ribosomes. The nucleus is expelled during the process of differentiation from an
1031:
6018:
5368:
Skutelsky E, Danon D (June 1970). "Comparative study of nuclear expulsion from the late erythroblast and cytokinesis". Primary.
3695:
Matera AG, Terns RM, Terns MP (March 2007). "Non-coding RNAs: lessons from the small nuclear and small nucleolar RNAs". Review.
412:, consisting of genes that are organized as heterochromatin only in certain cell types or at certain stages of development, and
7077:
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2187:
produced the same results for plants in 1884. This paved the way to assign the nucleus an important role in heredity. In 1873,
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108:
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speckles increases the kinetic efficiency of pre-mRNA splicing, ultimately boosting protein levels by modulation of splicing.
6977:
1177:
910:
encoding genes related to ribosomal function. The assembled ribosomal subunits are the largest structures passed through the
5249:"Mistargeting of B-type lamins at the end of mitosis: implications on cell survival and regulation of lamins A/C expression"
4024:"Clastosome: a subtype of nuclear body enriched in 19S and 20S proteasomes, ubiquitin, and protein substrates of proteasome"
7179:
3526:
2212:
2200:
1692:) and fungi, the centrosomes are intranuclear, and their nuclear envelope also does not disassemble during cell division.
405:, is the more compact form, and contains DNA that is infrequently transcribed. This structure is further categorized into
350:
322:
exist, made up of unique proteins, RNA molecules, and particular parts of the chromosomes. The best-known of these is the
5524:"Phylogenetic relationships between the Acantharea and the Polycystinea: a molecular perspective on Haeckel's Radiolaria"
4643:"LPS-induced down-regulation of signal regulatory protein {alpha} contributes to innate immune activation in macrophages"
3543:
Tripathi K, Parnaik VK (September 2008). "Differential dynamics of splicing factor SC35 during the cell cycle". Primary.
2589:
Barned S, Goodman AD, Mattson DH (February 1995). "Frequency of anti-nuclear antibodies in multiple sclerosis". Primary.
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5581:
Horton TR (2006). "The number of nuclei in basidiospores of 63 species of ectomycorrhizal
Homobasidiomycetes". Primary.
5438:
Hutter KJ, Stöhr M (1982). "Rapid detection of mutagen induced micronucleated erythrocytes by flow cytometry". Primary.
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Human red blood cells, like those of other mammals, lack nuclei. This occurs as a normal part of the cells' development.
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413:
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A review article about nuclear transport, explains the principles of the mechanism, and the various transport pathways
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1354:
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with discrete domains defined by dense localization of the transcription factor PTF, which promotes transcription of
362:
is stained blue. The distinct chromosome territories of chromosome 2 (red) and chromosome 9 (green) are stained with
6914:
Website covering structure and function of the nucleus from the
Department of Oncology at the University of Alberta.
7814:
3645:"Assembly of the nuclear transcription and processing machinery: Cajal bodies (coiled bodies) and transcriptosomes"
1370:
4744:"Stress induced nuclear granules form in response to accumulation of misfolded proteins in Caenorhabditis elegans"
4184:
Goebel HH, Warlo I (January 1997). "Nemaline myopathy with intranuclear rods--intranuclear rod myopathy". Review.
1433:
in DNA, helping it wind and unwind, as well as a large variety of transcription factors that regulate expression.
787:
Mutations in lamin genes leading to defects in filament assembly cause a group of rare genetic disorders known as
7898:
7652:
4986:
2103:
1905:
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in particular do not enter the nucleus. Now there is a body of evidence that under pathological conditions (e.g.
1137:
741:. The actual function of the veil is not clear, although it is excluded from the nucleolus and is present during
2634:"A systematic genomic screen implicates nucleocytoplasmic transport and membrane growth in nuclear size control"
1251:, body. Clastosomes are not typically present in normal cells, making them hard to detect. They form under high
7635:
7630:
7625:
7620:
7615:
7610:
7174:
1220:
210:
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de Roos AD (2006). "The origin of the eukaryotic cell based on conservation of existing interfaces". Primary.
1773:, which also lacks other organelles such as mitochondria, and serves primarily as a transport vessel to ferry
6959:
contains digitized commentaries and links to seminal research papers on the nucleus. Published online in the
3586:
Handwerger KE, Gall JG (January 2006). "Subnuclear organelles: new insights into form and function". Review.
898:. The transcription, post-transcriptional processing, and assembly of rRNA occurs in the nucleolus, aided by
2051:
1968:
1405:
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conditions within the nucleus and degrade once there is a decrease in activity or if cells are treated with
981:
955:
738:
7139:
6121:
López-García P, Moreira D (May 2006). "Selective forces for the origin of the eukaryotic nucleus". Review.
3092:"Nuclear lamins A and B1: different pathways of assembly during nuclear envelope formation in living cells"
2919:
Stuurman N, Heins S, Aebi U (1998). "Nuclear lamins: their structure, assembly, and interactions". Review.
2894:
Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P, eds. (2002). "Chapter 4: DNA and
Chromosomes".
32:
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4545:"Molecular dissection of nuclear paraspeckles: towards understanding the emerging world of the RNP milieu"
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382:, and during cell division the chromatin can be seen to form the well-defined chromosomes familiar from a
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5483:"The angiosperm phloem sieve tube system: a role in mediating traits important to modern agriculture"
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Boulikas T (1995). "Phosphorylation of transcription factors and control of the cell cycle". Review.
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2003:
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522:, from the rest of the cell. The size of the nucleus is correlated to the size of the cell, and this
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the molecule guanosine triphosphate (GTP) to release energy. The key GTPase in nuclear transport is
1563:
bound by exportins. The ability of importins and exportins to transport their cargo is regulated by
592:
membrane. As part of the endoplasmic reticulum membrane, the outer nuclear membrane is studded with
7928:
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240:, a double membrane that encloses the entire organelle and isolates its contents from the cellular
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Lamond AI, Spector DL (August 2003). "Nuclear speckles: a model for nuclear organelles". Review.
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2871:
2817:
2614:
2417:
2367:
2184:
2114:
1810:
1603:
1448:
884:
453:
441:
4496:"P54nrb forms a heterodimer with PSP1 that localizes to paraspeckles in an RNA-dependent manner"
2386:
1559:, which are bound by importins, while those transported from the nucleus to the cytoplasm carry
1468:. While in the nucleus, pre-mRNA is associated with a variety of proteins in complexes known as
749:, bind chromatin and disrupting their structure inhibits transcription of protein-coding genes.
6730:
6074:
Martin W (December 2005). "Archaebacteria (Archaea) and the origin of the eukaryotic nucleus".
2436:
2272:
Lodish H, Berk A, Matsudaira P, Kaiser CA, Krieger M, Scott MP, Zipursky SL, Darnell J (2004).
2121:" ("cell builder"). He believed that he had observed new cells assembling around "cytoblasts".
2046:
2027:
in modern poxviruses and eukaryotes. It has been suggested that the unresolved question of the
829:
7361:
7199:
7082:
6831:
6820:
A review article about the evolution of the nucleus, explaining a number of different theories
6799:
6763:
6710:
6674:
6622:
6579:
6515:
6467:
6432:
6389:
6332:
6281:
6232:
6173:
6138:
6091:
6054:
6050:
6044:
5999:
5950:
5893:
5873:
5855:
5796:
5747:
5698:
5647:
5598:
5563:
5504:
5455:
5420:
5385:
5348:
5313:
5278:
5229:
5180:
5142:
5093:
5040:
5009:
4965:
4940:
4917:
4882:
4839:
4802:
4775:
4724:
4672:
4623:
4574:
4525:
4443:
4391:
4342:
4307:
4258:
4201:
4166:
4117:
4063:
3997:
3940:
3905:
3849:
3809:
3769:
3712:
3674:
3603:
3560:
3510:
3460:
3414:
3365:
3308:
3248:
3209:
3170:
3121:
3072:
3062:
3054:
3035:
2985:
2936:
2899:
2863:
2803:
2780:
2729:
2665:
2606:
2571:
2518:
2489:"Active and inactive genes localize preferentially in the periphery of chromosome territories"
2469:
2409:
2359:
2324:
2281:
1893:
1510:
1489:
1166:
1149:
1016:
895:
888:
780:. Eight of these protofilaments form a lateral arrangement that is twisted to form a ropelike
299:
5005:
3805:
3141:"Alteration of nuclear lamin organization inhibits RNA polymerase II-dependent transcription"
2134:(1855) who decisively propagated the new paradigm that cells are generated solely by cells ("
564:(1). Other diagram labels show (2) the outer ring, (3) spokes, (4) basket, and (5) filaments.
7466:
7430:
6791:
6755:
6702:
6664:
6544:
6459:
6424:
6379:
6371:
6324:
6271:
6222:
6212:
6165:
6130:
6083:
5989:
5981:
5940:
5932:
5885:
5845:
5835:
5786:
5778:
5737:
5729:
5688:
5678:
5637:
5629:
5590:
5553:
5543:
5494:
5447:
5412:
5377:
5340:
5305:
5268:
5260:
5219:
5211:
5132:
5083:
5073:
5001:
4909:
4874:
4831:
4765:
4755:
4714:
4706:
4662:
4654:
4613:
4605:
4564:
4556:
4515:
4507:
4433:
4425:
4381:
4373:
4334:
4297:
4289:
4248:
4240:
4193:
4156:
4148:
4107:
4099:
4053:
4045:
3987:
3979:
3932:
3895:
3887:
3801:
3759:
3751:
3704:
3664:
3656:
3595:
3552:
3502:
3452:
3404:
3396:
3355:
3347:
3298:
3240:
3201:
3160:
3152:
3111:
3103:
3025:
2975:
2967:
2928:
2853:
2770:
2762:
2655:
2645:
2598:
2561:
2553:
2508:
2500:
2487:
Kurz A, Lampel S, Nickolenko JE, Bradl J, Benner A, Zirbel RM, et al. (December 1996).
2461:
2401:
2351:
2314:
2274:
2072:
2028:
1658:
1531:
1309:
1022:
Similar to Cajal bodies are Gemini of Cajal bodies, or gems, whose name is derived from the
876:
773:
679:
632:
628:
569:
561:
543:
515:
470:
427:. During interphase the chromatin organizes itself into discrete individual patches, called
307:
237:
6169:
4865:
Görlich D, Kutay U (1999). "Transport between the cell nucleus and the cytoplasm". Review.
1975:. Observations that myxobacteria are motile, can form multicellular complexes, and possess
1011:. CBs are involved in a number of different roles relating to RNA processing, specifically
623:. The nuclear pore complex is composed of approximately thirty different proteins known as
7669:
7458:
7062:
6967:
4469:
4022:
Lafarga M, Berciano MT, Pena E, Mayo I, Castaño JG, Bohmann D, et al. (August 2002).
2188:
1987:
1869:
1806:
1726:
1681:
1618:
1461:
1389:
1297:
1293:
1285:
1156:, and the rods themselves consist of mutant actin as well as other cytoskeletal proteins.
972:
663:
597:
402:
398:
284:
220:. Eukaryotic cells usually have a single nucleus, but a few cell types, such as mammalian
118:
5718:"Interleukin 4 induces cultured monocytes/macrophages to form giant multinucleated cells"
3385:"p53 mediates target gene association with nuclear speckles for amplified RNA expression"
2117:
proposed that the nucleus plays a role in generating cells, thus he introduced the name "
1629:
During its lifetime, a nucleus may be broken down or destroyed, either in the process of
6420:
6320:
6267:
6208:
5831:
5539:
5128:
3764:
3739:
3294:
2058:
7845:
7515:
7471:
7025:
6307:
Takemura M (May 2001). "Poxviruses and the origin of the eukaryotic nucleus". Primary.
5994:
5969:
5945:
5920:
5850:
5815:
5742:
5717:
5693:
5666:
5273:
5248:
5224:
5199:
5088:
5061:
4770:
4743:
4719:
4694:
4667:
4642:
4618:
4593:
4569:
4544:
4520:
4495:
4438:
4413:
4386:
4361:
4302:
4277:
4253:
4228:
4161:
4136:
4112:
4087:
3992:
3967:
3900:
3875:
3409:
3384:
3360:
3335:
3165:
3140:
3116:
3091:
2980:
2955:
2775:
2750:
2660:
2633:
2513:
2131:
2068:
2024:
2008:
1948:
1912:
1827:
1782:
1750:
1730:
1646:
1422:
1346:
1232:
1043:
921:, the nucleolus can be seen to consist of three distinguishable regions: the innermost
849:
824:
713:
703:
675:
511:
319:
245:
221:
217:
184:
6993:
6384:
6359:
6227:
6192:
5791:
5766:
5642:
5617:
5416:
5062:"The RanGTP Pathway: From Nucleo-Cytoplasmic Transport to Spindle Assembly and Beyond"
4197:
4058:
4023:
2566:
2541:
2526:
2019:". Another variant proposes that eukaryotes originated from early archaea infected by
302:
of molecules across the envelope. The pores cross both nuclear membranes, providing a
7912:
7308:
7268:
7243:
7159:
7072:
7047:
6706:
6599:
Brown R (1866). "On the Organs and Mode of
Fecundation of Orchidex and Asclepiadea".
6375:
6040:
6026:
5558:
5523:
5381:
4913:
4878:
4795:
3669:
3644:
3426:
2858:
2841:
2319:
2302:
2164:
2148:
2144:
2088:. Unlike mammalian red blood cells, those of other vertebrates still contain nuclei.
1960:
1822:
1696:
lamina. The destruction of the lamin networks is controlled by specialized apoptotic
1630:
1572:
1535:
1519:
1496:, and allows production of a large variety of proteins from a limited amount of DNA.
1426:
1264:
942:
907:
861:
857:
853:
789:
761:
612:
573:
483:
291:
233:
168:
134:
47:
6954:
6811:
6556:
6344:
6293:
5905:
5467:
5325:
5021:
4962:
Genome
Structure and Function: From Chromosomes Characterization to Genes Technology
4851:
4213:
3952:
3724:
3320:
3260:
2618:
615:
having around 20,000. The NPC provides selective transport of molecules between the
7699:
7479:
7288:
7260:
7238:
7231:
7211:
7151:
7067:
6693:
Görlich D, Kutay U (1999). "Transport between the cell nucleus and the cytoplasm".
6687:
A review article about nuclear lamins, explaining their structure and various roles
6535:
Cohen WD (1982). "The cytomorphic system of anucleate non-mammalian erythrocytes".
6479:
6025:. Washington, DC.: National Council for Science and the Environment. Archived from
5767:"Human giant cell tumors of bone identification and characterization of cell types"
5633:
5154:
4594:"Nuclear paraspeckles function in mediating gene regulatory and apoptotic pathways"
3821:
3572:
3522:
3472:
2724:
Lodish HF, Berk A, Kaiser C, Krieger M, Bretscher A, Ploegh H, et al. (2016).
2421:
2371:
2127:
2012:
1995:
1952:
1865:
1861:
1794:
1790:
1670:
1650:
1591:
1465:
1342:
994:
911:
879:
transcribes rDNA, which forms a large pre-rRNA precursor. This is cleaved into two
802:
644:
585:
557:
547:
530:
in animal cells. In human cells, the diameter of the nucleus is approximately six
387:
295:
124:
65:
6985:
6919:
6572:
4338:
3842:
2875:
1625:, stained light blue. All chromosomes but one are already at the metaphase plate.
196:
6795:
6759:
5889:
5840:
5683:
4229:"Compartmentalization within the nucleus: discovery of a novel subnuclear region"
3400:
2650:
2465:
2011:). One version of the proposal suggests that the nucleus evolved in concert with
848:
is the largest of the discrete densely stained, membraneless structures known as
7851:
7714:
7704:
7587:
7485:
7390:
7356:
7283:
7278:
7169:
6941:
contains peer-reviewed still images and video clips that illustrate the nucleus.
6905:
6877:
4710:
4609:
4429:
4377:
4276:
Pombo A, Cuello P, Schul W, Yoon JB, Roeder RG, Cook PR, Murphy S (March 1998).
4103:
3792:
Cioce M, Lamond AI (2005). "Cajal bodies: a long history of discovery". Review.
3351:
3228:
2227:
2122:
2092:
1956:
1843:
1786:
1770:
1714:
1689:
1662:
1607:
1480:
1259:. The scarcity of clastosomes in cells indicates that they are not required for
1252:
1189:
1142:
930:
765:
757:
678:, can diffuse through the cell membrane and into the cytoplasm, where they bind
671:
655:
624:
616:
519:
394:
303:
256:
190:
74:
44:
40:
17:
6463:
6428:
6360:"A hypothesis for DNA viruses as the origin of eukaryotic replication proteins"
6197:
Proceedings of the National Academy of Sciences of the United States of America
5985:
5594:
5528:
Proceedings of the National Academy of Sciences of the United States of America
3755:
3487:
1967:
host. The nuclear membrane may have served to protect the genome from damaging
1908:, sometimes accompany inflammation and are also implicated in tumor formation.
1376:
traditional view of moving replication forks along stagnant DNA, a concept of
552:
7792:
7605:
7385:
7325:
7318:
7303:
7298:
7293:
7248:
7194:
7184:
7134:
7129:
7119:
6156:
Fuerst JA (2005). "Intracellular compartmentation in planctomycetes". Review.
6109:
6087:
5936:
5309:
4760:
4293:
3983:
3599:
3556:
3506:
3303:
3278:
3244:
3205:
2355:
2176:
2152:
2064:
1901:
1896:
can also be abnormal in humans; for example, cells arising from the fusion of
1885:
1881:
1831:
1797:, which is the immediate precursor of the mature erythrocyte. The presence of
1746:
1666:
1622:
1568:
1322:
1314:
1289:
1260:
1204:
1200:
1003:
976:
742:
640:
608:
531:
437:
424:
375:
371:
355:
339:
260:
229:
163:
51:
5548:
5345:
Normal cell morphology in canine and feline cytology: an identification guide
5078:
4641:
Kong XN, Yan HX, Chen L, Dong LW, Yang W, Liu Q, et al. (October 2007).
3740:"Genome organization around nuclear speckles drives mRNA splicing efficiency"
2956:"Pathway of incorporation of microinjected lamin A into the nuclear envelope"
2704:
Alberts B, Johnson A, Lewis J, Morgan D, Raff M, Roberts K, Walter P (2015).
54:, thus their entire nuclei are labeled. On the left, a cell is going through
7745:
7694:
7597:
7406:
7368:
7341:
7313:
7221:
7206:
7189:
7124:
7097:
7029:
6252:"Viral eukaryogenesis: was the ancestor of the nucleus a complex DNA virus?"
5667:"Transposon Domestication versus Mutualism in Ciliate Genome Rearrangements"
4511:
4049:
3621:
2172:
2168:
1991:
1980:
1935:
1928:
1889:
1877:
1634:
1614:
1430:
1421:, which unwind the double-stranded DNA molecule to facilitate access to it,
1212:
934:
869:
845:
838:
818:
806:
527:
495:
491:
383:
379:
323:
241:
214:
152:
148:
92:
6803:
6782:
Pennisi E (August 2004). "Evolutionary biology. The birth of the nucleus".
6714:
6678:
6471:
6436:
6393:
6336:
6285:
6236:
6217:
6177:
6142:
6095:
6003:
5954:
5897:
5859:
5702:
5602:
5508:
5424:
5317:
5282:
5233:
5146:
5097:
5035:
Watson JD, Baker TA, Bell SP, Gann A, Levine M, Losick R (2004). "Ch9–10".
5013:
4921:
4886:
4843:
4779:
4728:
4676:
4627:
4578:
4529:
4447:
4395:
4346:
4170:
4121:
4067:
4001:
3813:
3773:
3716:
3678:
3660:
3607:
3564:
3514:
3464:
3418:
3369:
3312:
3252:
3213:
3174:
3125:
3107:
3076:
3039:
2954:
Goldman AE, Moir RD, Montag-Lowy M, Stewart M, Goldman RD (November 1992).
2932:
2867:
2784:
2766:
2669:
2632:
Kume K, Cantwell H, Neumann FR, Jones AW, Snijders AP, Nurse P (May 2017).
2504:
2363:
2328:
1927:
As the major defining characteristic of the eukaryotic cell, the nucleus's
1761:
1457:
1002:
A nucleus typically contains between one and ten compact structures called
674:, as well as other small lipid-soluble molecules involved in intercellular
6767:
6328:
6276:
6251:
5800:
5751:
5733:
5651:
5567:
5459:
5389:
5264:
5184:
4311:
4262:
4244:
4205:
4152:
3944:
3937:
10.1002/(sici)1097-4644(19980801)70:2<181::aid-jcb4>3.0.co;2-k
3909:
3156:
2989:
2971:
2940:
2610:
2575:
2522:
2473:
2413:
518:. The nuclear envelope separates the fluid inside the nucleus, called the
7674:
7351:
7107:
7092:
7087:
5499:
5482:
4658:
2602:
2222:
2180:
2096:
2020:
2016:
1943:
1897:
1835:
1697:
1642:
1552:
1548:
1425:, which bind to the DNA promoter to synthesize the growing RNA molecule,
1418:
1350:
1338:
1215:. During telophase, when the two daughter nuclei are formed, there is no
892:
880:
801:
in those with the condition. The exact mechanism by which the associated
794:
667:
593:
523:
475:
420:
374:. Each human cell contains roughly two meters of DNA. During most of the
327:
158:
102:
4560:
1983:
similar to eukarya, support a bacterial origin for the eukaryotic cell.
1515:
310:
by carrier proteins while allowing free movement of small molecules and
84:
7723:
7373:
7273:
7226:
7102:
6669:
6652:
6548:
6134:
5876:(August 2004). "Evolutionary biology. The birth of the nucleus". News.
5451:
5215:
4835:
3966:
Navascues J, Berciano MT, Tucker KE, Lafarga M, Matera AG (June 2004).
3923:
Matera AG (August 1998). "Of coiled bodies, gems, and salmon". Review.
3030:
3013:
2405:
1972:
1964:
1939:
1873:
1857:
1852:
1798:
1701:
1685:
1527:
1473:
1318:
1208:
753:
730:
620:
272:
268:
144:
139:
55:
6651:
Goldman RD, Gruenbaum Y, Moir RD, Shumaker DK, Spann TP (March 2002).
5782:
3968:"Targeting SMN to Cajal bodies and nuclear gems during neuritogenesis"
3012:
Goldman RD, Gruenbaum Y, Moir RD, Shumaker DK, Spann TP (March 2002).
2751:"Yeast nucleoporins involved in passive nuclear envelope permeability"
2557:
1998:
archaea and bacteria to generate the nucleus and the eukaryotic cell.
436:
Antibodies to certain types of chromatin organization, in particular,
7775:
7765:
7755:
7750:
7662:
7645:
7580:
7570:
7565:
7560:
7555:
7550:
7545:
7540:
7216:
3488:"Differential dynamics of splicing factor SC35 during the cell cycle"
2160:
2107:
2085:
1976:
1916:
1774:
1564:
1012:
1008:
903:
726:
397:
is the less compact DNA form, and contains genes that are frequently
252:
5137:
5112:
3708:
3456:
3189:
4900:
Hozák P, Cook PR (February 1994). "Replication factories". Review.
3891:
3139:
Spann TP, Goldman AE, Wang C, Huang S, Goldman RD (February 2002).
2842:"Mechanisms of receptor-mediated nuclear import and nuclear export"
1325:
is an enzyme responsible for the first step of glycolysis, forming
682:
proteins that are trafficked into the nucleus. There they serve as
7865:
7770:
7760:
7733:
7728:
7657:
7640:
7535:
7530:
7525:
7520:
7510:
7505:
7500:
7495:
7490:
3383:
Alexander KA, Coté A, Nguyen SC, Zhang L, Berger SL (March 2021).
2156:
2057:
2045:
1839:
1778:
1760:
1705:
1514:
1399:
1153:
1027:
993:
967:
868:(NOR). The main roles of the nucleolus are to synthesize rRNA and
798:
793:. The most notable laminopathy is the family of diseases known as
721:
636:
551:
469:
386:. A small fraction of the cell's genes are located instead in the
31:
2303:"Chromatin dynamics at DNA replication, transcription and repair"
760:
domain used by two monomers to coil around each other, forming a
7830:
7825:
7802:
7797:
7785:
7780:
7738:
4937:
Protein Synthesis and Ribosome Structure: Translating the Genome
1934:
The first model known as the "syntrophic model" proposes that a
1846:
have naturally multinucleated cells. Other examples include the
1674:
1599:
1596:
1484:
979:. B snurposomes are found in the amphibian oocyte nuclei and in
603:
In a mammalian nuclear envelope there are between 3000 and 4000
276:
36:
7426:
6997:
5814:
Imanian B, Pombert JF, Dorrell RG, Burki F, Keeling PJ (2012).
5665:
Vogt A, Goldman AD, Mochizuki K, Landweber LF (1 August 2013).
752:
Like the components of other intermediate filaments, the lamin
7346:
6960:
5765:
Goldring SR, Roelke MS, Petrison KK, Bhan AK (February 1987).
3690:
3688:
2749:
Shulga N, Mosammaparast N, Wozniak R, Goldfarb DS (May 2000).
1610:
1523:
1216:
959:
768:. Two of these dimer structures then join side by side, in an
690:; in the absence of a ligand, many such receptors function as
503:
487:
359:
315:
311:
264:
2898:(4th ed.). New York: Garland Science. pp. 191–234.
2191:
postulated the equivalence of the maternal and paternal germ
1951:
archaea, invaded and lived within bacteria similar to modern
954:
examined by electron microscopy they are seen as clusters of
248:, a network within the nucleus that adds mechanical support.
6193:"The origin of the eukaryotic cell: a genomic investigation"
2199:
were rediscovered at the beginning of the 20th century; the
1223:
so the protein components instead form a perinucleolar cap.
902:(snoRNA) molecules, some of which are derived from spliced
4693:
Carmo-Fonseca M, Berciano MT, Lafarga M (September 2010).
4137:"Paraspeckles: nuclear bodies built on long noncoding RNA"
875:
In the first step of ribosome assembly, a protein called
419:
that consists of chromosome structural components such as
6653:"Nuclear lamins: building blocks of nuclear architecture"
3014:"Nuclear lamins: building blocks of nuclear architecture"
2031:
could be related to the viral eukaryogenesis hypothesis.
6621:. Berlin, Heidelberg, New York, Tokyo: Springer Verlag.
2385:
Schardin M, Cremer T, Hager HD, Lang M (December 1985).
1621:
can be seen, stained green, attached to the two sets of
6971:
6935:. The American Society for Cell Biology. Archived from
5039:(5th ed.). Peason Benjamin Cummings; CSHL Press.
4987:"Mechanisms of alternative pre-messenger RNA splicing"
3090:
Moir RD, Yoon M, Khuon S, Goldman RD (December 2000).
1292:. The main function of the cell nucleus is to control
378:
these are organized in a DNA-protein complex known as
6049:. San Francisco: W. H. Freeman and Company. pp.
4688:
4686:
3190:"Aging and nuclear organization: lamins and progeria"
3055:"Dynamics of nuclear lamina assembly and disassembly"
1860:
have two kinds of nuclei in a single cell, a somatic
1345:
responses, transcription is induced in response to a
1152:. This condition typically results from mutations in
6978:"Cytoplasmic patterns generated by human antibodies"
4227:
Saunders WS, Cooke CA, Earnshaw WC (November 1991).
3643:
Gall JG, Bellini M, Wu Z, Murphy C (December 1999).
2050:
Oldest known depiction of cells and their nuclei by
1534:
across the nuclear membrane in a process called the
1476:
tail is only added after transcription is complete.
514:, and is enveloped in a double membrane called the
236:. The main structures making up the nucleus are the
27:
Eukaryotic membrane-bounded organelle containing DNA
7813:
7713:
7683:
7596:
7465:
7399:
7334:
7259:
7150:
7036:
1781:to the body's tissues. Erythrocytes mature through
1019:(snRNA) maturation, and histone mRNA modification.
314:. Movement of large molecules such as proteins and
251:The cell nucleus contains nearly all of the cell's
64:
6571:
4794:
3841:
2273:
2140:"). The function of the nucleus remained unclear.
2023:, on the basis of observed similarity between the
1479:RNA splicing, carried out by a complex called the
745:. Lamin structures that make up the veil, such as
452:(ANA) and have also been observed in concert with
7881:transcription factors and intracellular receptors
5347:. Hoboken, NJ: John Wiley & Sons. p. 6.
4543:Nakagawa S, Yamazaki T, Hirose T (October 2018).
3738:Bhat P, Chow A, Emert B, et al. (May 2024).
3272:
3270:
1349:such as that initiated by the signaling molecule
712:provide the nucleus with mechanical support: The
3059:Symposia of the Society for Experimental Biology
6922:. Medical Research Council Human Genetics Unit.
6695:Annual Review of Cell and Developmental Biology
5522:Zettler LA, Sogin ML, Caron DA (October 1997).
5113:"Cell biology: ripping up the nuclear envelope"
4867:Annual Review of Cell and Developmental Biology
4017:
4015:
4013:
4011:
3794:Annual Review of Cell and Developmental Biology
2699:
2095:in 1804 and in more detail in 1831 by Scottish
290:Because the nuclear envelope is impermeable to
5173:Critical Reviews in Eukaryotic Gene Expression
4360:Lallemand-Breitenbach V, de Thé H (May 2010).
2719:
2717:
2715:
2697:
2695:
2693:
2691:
2689:
2687:
2685:
2683:
2681:
2679:
1447:Newly synthesized mRNA molecules are known as
1341:-controlled genes, which are involved in most
837:of a cell nucleus, showing the darkly stained
502:The nucleus contains nearly all of the cell's
456:as part of general immune system dysfunction.
7438:
7009:
6947:"Nuclear Envelope and Nuclear Import Section"
5200:"The cell biology of open and closed mitosis"
4088:"Functional architecture in the cell nucleus"
3869:
3867:
3865:
3438:
3436:
1884:. Other multinucleate cells in the human are
1414:nucleus, mRNA produced needs to be exported.
1136:Beyond the nuclear bodies first described by
8:
6358:Villarreal LP, DeFilippis VR (August 2000).
5970:"Open Questions on the Origin of Eukaryotes"
4801:(3rd ed.). New York: Worth Publishers.
4494:Fox AH, Bond CS, Lamond AI (November 2005).
2835:
2833:
6110:https://doi.org/10.1007/978-3-319-65536-9_7
5968:López-García P, Moreira D (November 2015).
5166:
5164:
5066:Frontiers in Cell and Developmental Biology
4407:
4405:
3007:
3005:
3003:
3001:
2999:
2728:(Eighth ed.). New York: W.H. Freeman.
2135:
797:, which causes the appearance of premature
7687:
7445:
7431:
7423:
7016:
7002:
6994:
6619:Von der Zellenlehre zur Chromosomentheorie
6612:
6610:
4742:Sampuda KM, Riley M, Boyd L (April 2017).
4699:Cold Spring Harbor Perspectives in Biology
4468:. Nuclear Protein Database. Archived from
4418:Cold Spring Harbor Perspectives in Biology
4366:Cold Spring Harbor Perspectives in Biology
3876:"Coiled bodies and gems: Janus or gemini?"
3835:
3833:
3831:
3340:Cold Spring Harbor Perspectives in Biology
2822:: CS1 maint: location missing publisher (
2540:Rothfield NF, Stollar BD (November 1967).
1665:, which in turn are attached to different
1657:occurs by the replicated chromosomes, the
1030:(snRNPs), but do contain a protein called
611:only have about a few hundred, with large
6749:
6668:
6383:
6275:
6226:
6216:
5993:
5944:
5849:
5839:
5790:
5741:
5692:
5682:
5641:
5557:
5547:
5498:
5272:
5223:
5136:
5087:
5077:
4769:
4759:
4718:
4666:
4617:
4568:
4519:
4437:
4385:
4301:
4252:
4160:
4111:
4057:
4039:
3991:
3899:
3763:
3668:
3486:Tripathi K, Parnaik VK (September 2008).
3408:
3359:
3302:
3164:
3115:
3029:
2979:
2857:
2802:(Fifth ed.). New York. p. 242.
2774:
2659:
2649:
2565:
2512:
2455:
2318:
2267:
2265:
2263:
1470:heterogeneous ribonucleoprotein particles
720:The nuclear lamina is composed mostly of
275:, that protect and organize the DNA. The
50:. The central and rightmost cells are in
5919:Devos DP, Gräf R, Field MC (June 2014).
5006:10.1146/annurev.biochem.72.121801.161720
4793:Lehninger AL, Nelson DL, Cox MM (2000).
4489:
4487:
4459:
4457:
4081:
4079:
4077:
3806:10.1146/annurev.cellbio.20.010403.103738
3787:
3785:
3783:
2708:(6 ed.). New York: Garland Science.
2261:
2259:
2257:
2255:
2253:
2251:
2249:
2247:
2245:
2243:
1590:
1329:from glucose. At high concentrations of
1280:that is segregated from the location of
1047:
828:
440:, have been associated with a number of
349:
6731:"Structure and function in the nucleus"
5716:McInnes A, Rennick DM (February 1988).
3334:Spector DL, Lamond AI (February 2011).
3229:"Nucleolus: from structure to dynamics"
2889:
2887:
2885:
2840:Pemberton LF, Paschal BM (March 2005).
2437:"Structure and function in the nucleus"
2239:
2001:The most controversial model, known as
1317:, a cellular pathway for breaking down
306:through which larger molecules must be
6191:Hartman H, Fedorov A (February 2002).
6170:10.1146/annurev.micro.59.030804.121258
3697:Nature Reviews. Molecular Cell Biology
3532:from the original on 15 November 2011.
3445:Nature Reviews. Molecular Cell Biology
3277:Lamond AI, Sleeman JE (October 2003).
2815:
2280:(5th ed.). New York: WH Freeman.
2155:eggs, showing that the nucleus of the
1725:Initially, it has been suspected that
852:found in the nucleus. It forms around
61:
6972:The American Society for Cell Biology
6729:Lamond AI, Earnshaw WC (April 1998).
6021:. In Monosson E, Cleveland C (eds.).
5771:The Journal of Clinical Investigation
2546:The Journal of Clinical Investigation
2435:Lamond AI, Earnshaw WC (April 1998).
1284:in the cytoplasm, allowing levels of
506:, surrounded by a network of fibrous
225:
89:Components of a typical animal cell:
7:
6578:. New Haven: Yale University Press.
5722:The Journal of Experimental Medicine
5111:Lippincott-Schwartz J (March 2002).
4797:Lehninger principles of biochemistry
4647:The Journal of Experimental Medicine
4466:"Nuclear Compartments: Paraspeckles"
3622:"Cellular component Nucleus speckle"
3188:Mounkes LC, Stewart CL (June 2004).
1830:cells contain multiple nuclei. Most
937:), which in turn is bordered by the
627:. The pores are about 60–80 million
4592:Pisani G, Baron B (December 2019).
3279:"Nuclear substructure and dynamics"
3227:Hernandez-Verdun D (January 2006).
474:Diagram of the nucleus showing the
202:
6924:Information on nuclear components.
4135:Bond CS, Fox AH (September 2009).
3880:American Journal of Human Genetics
3874:Matera AG, Frey MR (August 1998).
2301:Ehrenhofer-Murray AE (June 2004).
2091:The nucleus was also described by
1856:, which have two nuclei per cell.
1203:, paraspeckles are present during
639:) to several hundred proteins (in
393:There are two types of chromatin.
25:
5974:Trends in Ecology & Evolution
5247:Steen RL, Collas P (April 2001).
2147:published several studies on the
1453:post-transcriptional modification
1443:Post-transcriptional modification
1067:
864:(rRNA). These regions are called
708:In animal cells, two networks of
364:fluorescent in situ hybridization
259:is often organized into multiple
6826:Pollard TD, Earnshaw WC (2004).
6707:10.1146/annurev.cellbio.15.1.607
6376:10.1128/JVI.74.15.7079-7084.2000
4879:10.1146/annurev.cellbio.15.1.607
4824:Biochemical Society Transactions
4086:Dundr M, Misteli T (June 2001).
3925:Journal of Cellular Biochemistry
3840:Pollard TD, Earnshaw WC (2004).
3053:Broers JL, Ramaekers FC (2004).
2859:10.1111/j.1600-0854.2005.00270.x
2320:10.1111/j.1432-1033.2004.04162.x
2307:European Journal of Biochemistry
1753:, or from faulty cell division.
1276:The nucleus provides a site for
1028:small nuclear ribonucleoproteins
941:(GC) (that contains the protein
460:Nuclear structures and landmarks
83:
6951:Landmark Papers in Cell Biology
6601:Miscellaneous Botanical Works I
5925:Current Opinion in Cell Biology
5481:Ham BK, Lucas WJ (April 2014).
4935:Nierhaus KH, Wilson DN (2004).
4412:Fox AH, Lamond AI (July 2010).
3233:Histochemistry and Cell Biology
3194:Current Opinion in Cell Biology
1451:or pre-mRNA. They must undergo
1054:
6920:"The Nuclear Protein Database"
6906:"Learn about the Cell Nucleus"
6510:. Frankfurt am Main, Germany:
6409:Journal of Theoretical Biology
6309:Journal of Molecular Evolution
6256:Journal of Molecular Evolution
5634:10.1128/MMBR.55.4.706-732.1991
5487:Journal of Experimental Botany
5198:Boettcher B, Barral Y (2013).
2075:in 1882. The nucleus contains
1178:Promyelocytic leukemia protein
805:changes give rise to the aged
694:that repress gene expression.
326:, involved in the assembly of
1:
7180:Microtubule organizing center
6945:Gall JG, McIntosh JR (eds.).
6158:Annual Review of Microbiology
5417:10.1016/S1383-5718(99)00216-8
5037:Molecular Biology of the Gene
4994:Annual Review of Biochemistry
4500:Molecular Biology of the Cell
4339:10.1016/j.cellsig.2004.03.020
4198:10.1016/S0960-8966(96)00404-X
4028:Molecular Biology of the Cell
3649:Molecular Biology of the Cell
2921:Journal of Structural Biology
2896:Molecular Biology of the Cell
2706:Molecular Biology of the Cell
2213:Computed Corpuscle Sectioning
2201:chromosome theory of heredity
1737:) IgG can enter the nucleus.
1429:, which change the amount of
1060:
635:and consist of around 50 (in
401:by the cell. The other type,
279:within these chromosomes are
228:, and a few others including
6883:Resources in other libraries
6796:10.1126/science.305.5685.766
6760:10.1126/science.280.5363.547
5890:10.1126/science.305.5685.766
5841:10.1371/journal.pone.0043763
5684:10.1371/journal.pgen.1003659
5618:"The biology of Giardia spp"
5382:10.1016/0014-4827(70)90536-7
5060:Cavazza T, Vernos I (2015).
4914:10.1016/0962-8924(94)90009-4
3401:10.1016/j.molcel.2021.03.006
3395:(8): S1097-2765(21)00174-X.
2651:10.1371/journal.pgen.1006767
2466:10.1126/science.280.5363.547
2167:'s theory that the complete
2034:A more recent proposal, the
1557:nuclear localization signals
1337:For example, in the case of
446:systemic lupus erythematosus
130:Smooth endoplasmic reticulum
6046:Symbiosis in Cell Evolution
5253:The Journal of Cell Biology
4711:10.1101/cshperspect.a000703
4610:10.1016/j.ncrna.2019.11.002
4430:10.1101/cshperspect.a000687
4378:10.1101/cshperspect.a000661
4233:The Journal of Cell Biology
4141:The Journal of Cell Biology
3352:10.1101/cshperspect.a000646
3145:The Journal of Cell Biology
3096:The Journal of Cell Biology
2960:The Journal of Cell Biology
2755:The Journal of Cell Biology
2493:The Journal of Cell Biology
2015:to form an early cellular "
1809:, this condition occurs in
1492:. This process is known as
1355:nuclear localisation signal
866:nucleolar organizer regions
114:Rough endoplasmic reticulum
7945:
7815:Transition nuclear protein
6830:. Philadelphia: Saunders.
6508:Geschichte der Mikroskopie
6464:10.1162/artl.2006.12.4.513
6429:10.1016/j.jtbi.2006.05.015
6250:Bell PJ (September 2001).
5986:10.1016/j.tree.2015.09.005
5921:"Evolution of the nucleus"
5595:10.3852/mycologia.98.2.233
5370:Experimental Cell Research
5210:(3). Austin, Tex.: 160–5.
4464:Fox A, Bickmore W (2004).
3848:. Philadelphia: Saunders.
3756:10.1038/s41586-024-07429-6
1906:giant multinucleated cells
1820:
1508:
1440:
1393:
1387:
1371:Eukaryotic DNA replication
1368:
1288:that are not available to
1187:
1050:Subnuclear structure sizes
1041:
822:
816:
701:
541:
538:Nuclear envelope and pores
463:
343:
337:
58:and its DNA has condensed.
7894:
7874:
7861:
7841:
7690:
7653:Perinucleolar compartment
6961:Image & Video Library
6933:Image & Video Library
6878:Resources in your library
6088:10.1016/j.mib.2005.10.004
5937:10.1016/j.ceb.2014.01.004
5616:Adam RD (December 1991).
5310:10.1007/s10067-007-0597-y
4761:10.1186/s12860-017-0136-x
4104:10.1042/0264-6021:3560297
3984:10.1007/s00412-004-0285-5
3600:10.1016/j.tcb.2005.11.005
3557:10.1007/s12038-008-0054-3
3507:10.1007/s12038-008-0054-3
3304:10.1016/j.cub.2003.10.012
3245:10.1007/s00418-005-0046-4
3206:10.1016/j.ceb.2004.03.009
2356:10.1007/s10577-005-1021-6
2203:was therefore developed.
2104:Linnean Society of London
1938:relationship between the
1551:to enter the nucleus and
1304:Cell compartmentalization
1065:
1055:
927:dense fibrillar component
925:(FCs), surrounded by the
298:are required to regulate
82:
73:
7175:Prokaryotic cytoskeleton
6929:"The Nucleus Collection"
5549:10.1073/pnas.94.21.11411
5079:10.3389/fcell.2015.00082
1721:Disease-related dynamics
1587:Assembly and disassembly
1542:nuclear transport cycle.
1032:survival of motor neuron
917:When observed under the
809:is not well understood.
737:, that is visible using
211:membrane-bound organelle
151:; with which, comprises
6657:Genes & Development
5622:Microbiological Reviews
4695:"Orphan nuclear bodies"
4598:Non-Coding RNA Research
4512:10.1091/mbc.E05-06-0587
4294:10.1093/emboj/17.6.1768
4186:Neuromuscular Disorders
4092:The Biochemical Journal
4050:10.1091/mbc.e02-03-0122
3018:Genes & Development
2798:Alberts, Bruce (2019).
2143:Between 1877 and 1878,
2137:Omnis cellula e cellula
2052:Antonie van Leeuwenhoek
1969:reactive oxygen species
1633:or as a consequence of
1500:Dynamics and regulation
1396:Transcription factories
1300:during the cell cycle.
982:Drosophila melanogaster
956:interchromatin granules
772:arrangement, to form a
739:fluorescence microscopy
450:anti-nuclear antibodies
6218:10.1073/pnas.032658599
5341:"Nuclear Morphologies"
4902:Trends in Cell Biology
3661:10.1091/mbc.10.12.4385
3588:Trends in Cell Biology
3545:Journal of Biosciences
3495:Journal of Biosciences
3108:10.1083/jcb.151.6.1155
2933:10.1006/jsbi.1998.3987
2800:Essential cell biology
2767:10.1083/jcb.149.5.1027
2726:Molecular Cell Biology
2505:10.1083/jcb.135.5.1195
2276:Molecular Cell Biology
2218:Nucleus (neuroanatomy)
2136:
2080:
2055:
2036:exomembrane hypothesis
1766:
1626:
1561:nuclear export signals
1543:
1437:Processing of pre-mRNA
1410:
1138:Santiago Ramón y Cajal
999:
841:
710:intermediate filaments
605:nuclear pore complexes
582:outer nuclear membrane
565:
560:on the surface of the
508:intermediate filaments
499:
480:outer nuclear membrane
430:chromosome territories
367:
201: 'kernel, seed';
195:
189:
59:
7165:Intermediate filament
7058:Endoplasmic reticulum
6574:The Birth of the Cell
6329:10.1007/s002390010171
6277:10.1007/s002390010215
6023:Encyclopedia of Earth
5734:10.1084/jem.167.2.598
5298:Clinical Rheumatology
5265:10.1083/jcb.153.3.621
4245:10.1083/jcb.115.4.919
4153:10.1083/jcb.200906113
3157:10.1083/jcb.200112047
2972:10.1083/jcb.119.4.725
2529:on 29 September 2007.
2106:. Brown was studying
2061:
2049:
1892:. Multinucleated and
1870:skeletal muscle cells
1764:
1639:programmed cell death
1594:
1518:
1406:transcription factory
1403:
1378:replication factories
1278:genetic transcription
1257:proteasome inhibitors
1227:Perichromatin fibrils
997:
990:Cajal bodies and gems
929:(DFC) (that contains
832:
823:Further information:
684:transcription factors
590:endoplasmic reticulum
556:A cross section of a
555:
473:
464:Further information:
448:. These are known as
353:
344:Further information:
147:(fluid that contains
35:
7412:Extracellular matrix
6982:AntibodyPatterns.com
6966:10 June 2011 at the
6957:on 17 November 2006.
6939:on 12 November 2006.
6512:Verlag Harri Deutsch
4960:Nicolini CA (1997).
4659:10.1084/jem.20062611
4472:on 10 September 2008
4362:"PML nuclear bodies"
3624:. UniProt: UniProtKB
2603:10.1212/WNL.45.2.384
2077:polytene chromosomes
2004:viral eukaryogenesis
1994:, evolved first and
1848:intestinal parasites
1817:Multinucleated cells
1532:actively transported
1494:alternative splicing
1331:fructose-6-phosphate
1160:PIKA and PTF domains
1038:Other nuclear bodies
1024:Gemini constellation
692:histone deacetylases
686:when bound to their
346:Nuclear organization
308:actively transported
267:dotted with various
7606:Cajal (coiled) body
7467:Envelope (membrane)
7115:Cytoplasmic granule
6493:Van Leeuwenhoek A.
6421:2006JThBi.243...54B
6364:Journal of Virology
6321:2001JMolE..52..419T
6268:2001JMolE..53..251L
6209:2002PNAS...99.1420H
6076:Curr Opin Microbiol
5832:2012PLoSO...743763I
5540:1997PNAS...9411411A
5129:2002Natur.416...31L
4561:10.1098/rsob.180150
4327:Cellular Signalling
3295:2003CBio...13.R825L
2344:Chromosome Research
1811:sieve tube elements
1735:lupus erythematosus
1675:CDC2 protein kinase
1449:primary transcripts
1327:glucose-6-phosphate
1321:to produce energy.
1052:
1013:small nucleolar RNA
919:electron microscope
900:small nucleolar RNA
881:large rRNA subunits
835:electron micrograph
764:structure called a
466:Nuclear equivalence
442:autoimmune diseases
105:(dots as part of 5)
76:Animal cell diagram
7453:Structures of the
7140:Weibel–Palade body
7024:Structures of the
6988:on 2 January 2007.
6670:10.1101/gad.960502
6549:10.1007/BF01283036
6506:Gerlach D (2009).
6135:10.1002/bies.20413
5500:10.1093/jxb/ert417
5452:10.1007/bf00496738
5216:10.4161/nucl.24676
4836:10.1042/BST0330265
3336:"Nuclear speckles"
3031:10.1101/gad.960502
2406:10.1007/BF00388452
2185:Eduard Strasburger
2115:Matthias Schleiden
2081:
2071:cell published by
2056:
1767:
1627:
1544:
1411:
1298:replication of DNA
1207:and during all of
1173:PML-nuclear bodies
1062:Structure diameter
1048:
1000:
939:granular component
893:small rRNA subunit
870:assemble ribosomes
842:
735:nucleoplasmic veil
566:
500:
454:multiple sclerosis
368:
263:– long strands of
60:
7906:
7905:
7890:
7889:
7420:
7419:
7200:Spindle pole body
6859:Library resources
6837:978-0-7216-3360-2
6628:978-3-540-13987-4
6617:Cremer T (1985).
6585:978-0-300-07384-3
6570:Harris H (1999).
6521:978-3-8171-1781-9
6502:] (in Latin).
6060:978-0-7167-1256-5
6017:Hogan CM (2010).
5783:10.1172/JCI112838
5405:Mutation Research
5354:978-1-119-27891-7
5339:Ressel L (2017).
5046:978-0-8053-9603-4
4985:Black DL (2003).
4971:978-0-7923-4565-7
4946:978-3-527-30638-1
4808:978-1-57259-931-4
4098:(Pt 2): 297–310.
3855:978-0-7216-3360-2
2905:978-0-8153-4072-0
2735:978-1-4641-8339-3
2558:10.1172/JCI105669
2287:978-0-7167-2672-2
2102:in a talk at the
1894:binucleated cells
1659:sister chromatids
1511:Nuclear transport
1505:Nuclear transport
1490:protein sequences
1167:small nuclear RNA
1150:nemaline myopathy
1134:
1133:
1017:small nuclear RNA
949:Splicing speckles
923:fibrillar centers
860:, DNA coding for
358:nucleus in which
300:nuclear transport
177:
176:
16:(Redirected from
7936:
7899:nucleus diseases
7848:(Nucleoskeleton)
7688:
7447:
7440:
7433:
7424:
7018:
7011:
7004:
6995:
6989:
6984:. Archived from
6958:
6953:. Archived from
6940:
6923:
6913:
6901:
6841:
6815:
6771:
6753:
6744:(5363): 547–53.
6735:
6718:
6682:
6672:
6637:
6632:
6614:
6605:
6604:
6596:
6590:
6589:
6577:
6567:
6561:
6560:
6532:
6526:
6525:
6503:
6490:
6484:
6483:
6447:
6441:
6440:
6404:
6398:
6397:
6387:
6355:
6349:
6348:
6304:
6298:
6297:
6279:
6247:
6241:
6240:
6230:
6220:
6188:
6182:
6181:
6153:
6147:
6146:
6118:
6112:
6106:
6100:
6099:
6071:
6065:
6064:
6037:
6031:
6030:
6014:
6008:
6007:
5997:
5965:
5959:
5958:
5948:
5916:
5910:
5909:
5870:
5864:
5863:
5853:
5843:
5811:
5805:
5804:
5794:
5762:
5756:
5755:
5745:
5713:
5707:
5706:
5696:
5686:
5662:
5656:
5655:
5645:
5613:
5607:
5606:
5578:
5572:
5571:
5561:
5551:
5519:
5513:
5512:
5502:
5478:
5472:
5471:
5435:
5429:
5428:
5400:
5394:
5393:
5365:
5359:
5358:
5336:
5330:
5329:
5293:
5287:
5286:
5276:
5244:
5238:
5237:
5227:
5195:
5189:
5188:
5168:
5159:
5158:
5140:
5108:
5102:
5101:
5091:
5081:
5057:
5051:
5050:
5032:
5026:
5025:
4991:
4982:
4976:
4975:
4957:
4951:
4950:
4932:
4926:
4925:
4897:
4891:
4890:
4862:
4856:
4855:
4819:
4813:
4812:
4800:
4790:
4784:
4783:
4773:
4763:
4748:BMC Cell Biology
4739:
4733:
4732:
4722:
4690:
4681:
4680:
4670:
4638:
4632:
4631:
4621:
4589:
4583:
4582:
4572:
4540:
4534:
4533:
4523:
4491:
4482:
4481:
4479:
4477:
4461:
4452:
4451:
4441:
4409:
4400:
4399:
4389:
4357:
4351:
4350:
4333:(10): 1085–104.
4322:
4316:
4315:
4305:
4282:The EMBO Journal
4273:
4267:
4266:
4256:
4224:
4218:
4217:
4181:
4175:
4174:
4164:
4132:
4126:
4125:
4115:
4083:
4072:
4071:
4061:
4043:
4019:
4006:
4005:
3995:
3963:
3957:
3956:
3920:
3914:
3913:
3903:
3871:
3860:
3859:
3847:
3837:
3826:
3825:
3789:
3778:
3777:
3767:
3750:(5): 1165–1173.
3735:
3729:
3728:
3692:
3683:
3682:
3672:
3655:(12): 4385–402.
3640:
3634:
3633:
3631:
3629:
3618:
3612:
3611:
3583:
3577:
3576:
3540:
3534:
3533:
3531:
3492:
3483:
3477:
3476:
3440:
3431:
3430:
3412:
3380:
3374:
3373:
3363:
3331:
3325:
3324:
3306:
3274:
3265:
3264:
3224:
3218:
3217:
3185:
3179:
3178:
3168:
3136:
3130:
3129:
3119:
3087:
3081:
3080:
3050:
3044:
3043:
3033:
3009:
2994:
2993:
2983:
2951:
2945:
2944:
2916:
2910:
2909:
2891:
2880:
2879:
2861:
2837:
2828:
2827:
2821:
2813:
2795:
2789:
2788:
2778:
2746:
2740:
2739:
2721:
2710:
2709:
2701:
2674:
2673:
2663:
2653:
2629:
2623:
2622:
2586:
2580:
2579:
2569:
2537:
2531:
2530:
2525:. Archived from
2516:
2484:
2478:
2477:
2459:
2450:(5363): 547–53.
2441:
2432:
2426:
2425:
2391:
2382:
2376:
2375:
2339:
2333:
2332:
2322:
2298:
2292:
2291:
2279:
2269:
2139:
2073:Walther Flemming
2029:evolution of sex
1807:flowering plants
1757:Anucleated cells
1637:(the process of
1310:nuclear envelope
1296:and mediate the
1088:0.2–0.5 μm
1070:
1069:
1053:
877:RNA polymerase I
680:nuclear receptor
664:Steroid hormones
660:adaptor proteins
633:molecular weight
584:, perforated by
572:consists of two
570:nuclear envelope
562:nuclear envelope
544:Nuclear envelope
516:nuclear envelope
238:nuclear envelope
204:
121:(or, Golgi body)
87:
77:
68:
62:
21:
18:Nuclear proteins
7944:
7943:
7939:
7938:
7937:
7935:
7934:
7933:
7909:
7908:
7907:
7902:
7886:
7870:
7857:
7837:
7809:
7709:
7679:
7592:
7470:
7461:
7459:nuclear protein
7451:
7421:
7416:
7395:
7330:
7255:
7146:
7063:Golgi apparatus
7039:
7032:
7022:
6992:
6976:
6968:Wayback Machine
6944:
6927:
6917:
6910:cellnucleus.com
6904:
6892:
6889:
6888:
6887:
6867:
6866:
6862:
6854:
6849:
6838:
6825:
6790:(5685): 766–8.
6781:
6751:10.1.1.323.5543
6733:
6728:
6692:
6650:
6646:
6644:Further reading
6641:
6640:
6633:Online Version
6629:
6616:
6615:
6608:
6598:
6597:
6593:
6586:
6569:
6568:
6564:
6534:
6533:
6529:
6522:
6505:
6492:
6491:
6487:
6452:Artificial Life
6449:
6448:
6444:
6406:
6405:
6401:
6370:(15): 7079–84.
6357:
6356:
6352:
6306:
6305:
6301:
6249:
6248:
6244:
6190:
6189:
6185:
6155:
6154:
6150:
6120:
6119:
6115:
6107:
6103:
6073:
6072:
6068:
6061:
6039:
6038:
6034:
6029:on 11 May 2011.
6016:
6015:
6011:
5980:(11): 697–708.
5967:
5966:
5962:
5918:
5917:
5913:
5884:(5685): 766–8.
5872:
5871:
5867:
5813:
5812:
5808:
5764:
5763:
5759:
5715:
5714:
5710:
5677:(8): e1003659.
5664:
5663:
5659:
5615:
5614:
5610:
5580:
5579:
5575:
5534:(21): 11411–6.
5521:
5520:
5516:
5493:(7): 1799–816.
5480:
5479:
5475:
5437:
5436:
5432:
5402:
5401:
5397:
5367:
5366:
5362:
5355:
5338:
5337:
5333:
5304:(11): 1877–82.
5295:
5294:
5290:
5246:
5245:
5241:
5197:
5196:
5192:
5170:
5169:
5162:
5138:10.1038/416031a
5110:
5109:
5105:
5059:
5058:
5054:
5047:
5034:
5033:
5029:
4989:
4984:
4983:
4979:
4972:
4959:
4958:
4954:
4947:
4934:
4933:
4929:
4899:
4898:
4894:
4864:
4863:
4859:
4830:(Pt 1): 265–8.
4821:
4820:
4816:
4809:
4792:
4791:
4787:
4741:
4740:
4736:
4692:
4691:
4684:
4653:(11): 2719–31.
4640:
4639:
4635:
4591:
4590:
4586:
4542:
4541:
4537:
4506:(11): 5304–15.
4493:
4492:
4485:
4475:
4473:
4463:
4462:
4455:
4411:
4410:
4403:
4359:
4358:
4354:
4324:
4323:
4319:
4275:
4274:
4270:
4226:
4225:
4221:
4183:
4182:
4178:
4134:
4133:
4129:
4085:
4084:
4075:
4041:10.1.1.321.6138
4021:
4020:
4009:
3965:
3964:
3960:
3922:
3921:
3917:
3873:
3872:
3863:
3856:
3839:
3838:
3829:
3791:
3790:
3781:
3737:
3736:
3732:
3709:10.1038/nrm2124
3694:
3693:
3686:
3642:
3641:
3637:
3627:
3625:
3620:
3619:
3615:
3585:
3584:
3580:
3542:
3541:
3537:
3529:
3490:
3485:
3484:
3480:
3457:10.1038/nrm1172
3442:
3441:
3434:
3382:
3381:
3377:
3333:
3332:
3328:
3283:Current Biology
3276:
3275:
3268:
3239:(1–2): 127–37.
3226:
3225:
3221:
3187:
3186:
3182:
3138:
3137:
3133:
3089:
3088:
3084:
3069:
3052:
3051:
3047:
3011:
3010:
2997:
2953:
2952:
2948:
2918:
2917:
2913:
2906:
2893:
2892:
2883:
2839:
2838:
2831:
2814:
2810:
2797:
2796:
2792:
2748:
2747:
2743:
2736:
2723:
2722:
2713:
2703:
2702:
2677:
2644:(5): e1006767.
2631:
2630:
2626:
2588:
2587:
2583:
2552:(11): 1785–94.
2539:
2538:
2534:
2499:(5): 1195–205.
2486:
2485:
2481:
2457:10.1.1.323.5543
2439:
2434:
2433:
2429:
2389:
2384:
2383:
2379:
2341:
2340:
2336:
2313:(12): 2335–49.
2300:
2299:
2295:
2288:
2271:
2270:
2241:
2236:
2209:
2197:Mendelian rules
2189:August Weismann
2044:
2025:DNA polymerases
2009:viral envelopes
1988:Planctomycetota
1965:archaebacterial
1925:
1913:dinoflagellates
1864:and a germline
1825:
1819:
1759:
1751:red blood cells
1743:
1741:Nuclei per cell
1729:in general and
1727:immunoglobulins
1723:
1682:dinoflagellates
1661:, attaching to
1619:mitotic spindle
1589:
1567:, enzymes that
1513:
1507:
1502:
1462:polyadenylation
1445:
1439:
1423:RNA polymerases
1398:
1392:
1390:Gene expression
1386:
1384:Gene expression
1373:
1367:
1306:
1294:gene expression
1286:gene regulation
1274:
1241:
1229:
1192:
1186:
1175:
1162:
1118:0.5–1.0 μm
1108:0.2–1.0 μm
1078:0.2–2.0 μm
1066:
1046:
1040:
992:
973:phosphorylation
951:
827:
821:
815:
706:
700:
550:
542:Main articles:
540:
468:
462:
417:heterochromatin
410:heterochromatin
403:heterochromatin
348:
342:
336:
292:large molecules
285:gene expression
222:red blood cells
173:
119:Golgi apparatus
75:
66:
28:
23:
22:
15:
12:
11:
5:
7942:
7940:
7932:
7931:
7926:
7921:
7911:
7910:
7904:
7903:
7895:
7892:
7891:
7888:
7887:
7885:
7884:
7875:
7872:
7871:
7869:
7868:
7862:
7859:
7858:
7856:
7855:
7849:
7846:Nuclear matrix
7842:
7839:
7838:
7836:
7835:
7834:
7833:
7828:
7820:
7818:
7811:
7810:
7808:
7807:
7806:
7805:
7800:
7790:
7789:
7788:
7783:
7778:
7773:
7768:
7763:
7758:
7753:
7743:
7742:
7741:
7736:
7731:
7720:
7718:
7711:
7710:
7708:
7707:
7702:
7700:Dot (PML body)
7697:
7691:
7685:
7681:
7680:
7678:
7677:
7672:
7667:
7666:
7665:
7660:
7650:
7649:
7648:
7643:
7638:
7633:
7628:
7623:
7618:
7613:
7602:
7600:
7594:
7593:
7591:
7590:
7585:
7584:
7583:
7578:
7573:
7568:
7563:
7558:
7553:
7548:
7543:
7538:
7533:
7528:
7523:
7518:
7513:
7508:
7503:
7498:
7493:
7483:
7476:
7474:
7472:nuclear lamina
7463:
7462:
7452:
7450:
7449:
7442:
7435:
7427:
7418:
7417:
7415:
7414:
7409:
7403:
7401:
7397:
7396:
7394:
7393:
7388:
7383:
7382:
7381:
7376:
7366:
7365:
7364:
7359:
7354:
7344:
7338:
7336:
7335:Other internal
7332:
7331:
7329:
7328:
7323:
7322:
7321:
7316:
7311:
7306:
7301:
7296:
7291:
7286:
7281:
7271:
7265:
7263:
7257:
7256:
7254:
7253:
7252:
7251:
7246:
7236:
7235:
7234:
7229:
7224:
7219:
7209:
7204:
7203:
7202:
7197:
7192:
7187:
7177:
7172:
7167:
7162:
7156:
7154:
7148:
7147:
7145:
7144:
7143:
7142:
7137:
7132:
7127:
7122:
7112:
7111:
7110:
7105:
7100:
7095:
7090:
7085:
7075:
7070:
7065:
7060:
7055:
7050:
7044:
7042:
7034:
7033:
7023:
7021:
7020:
7013:
7006:
6998:
6991:
6990:
6974:
6942:
6925:
6915:
6902:
6886:
6885:
6880:
6875:
6869:
6868:
6857:
6856:
6855:
6853:
6852:External links
6850:
6848:
6847:
6843:
6842:
6836:
6822:
6821:
6817:
6816:
6778:
6777:
6773:
6772:
6725:
6724:
6720:
6719:
6689:
6688:
6684:
6683:
6647:
6645:
6642:
6639:
6638:
6627:
6606:
6591:
6584:
6562:
6527:
6520:
6485:
6442:
6399:
6350:
6299:
6242:
6183:
6148:
6113:
6101:
6066:
6059:
6032:
6009:
5960:
5911:
5865:
5806:
5757:
5728:(2): 598–611.
5708:
5657:
5608:
5573:
5514:
5473:
5440:Histochemistry
5430:
5395:
5360:
5353:
5331:
5288:
5239:
5190:
5160:
5123:(6876): 31–2.
5115:. Commentary.
5103:
5052:
5045:
5027:
5000:(1): 291–336.
4977:
4970:
4952:
4945:
4927:
4892:
4857:
4814:
4807:
4785:
4734:
4705:(9): a000703.
4682:
4633:
4604:(4): 128–134.
4584:
4555:(10): 180150.
4535:
4483:
4453:
4424:(7): a000687.
4414:"Paraspeckles"
4401:
4372:(5): a000661.
4352:
4317:
4288:(6): 1768–78.
4268:
4219:
4176:
4127:
4073:
4034:(8): 2771–82.
4007:
3978:(8): 398–409.
3958:
3915:
3892:10.1086/301992
3861:
3854:
3827:
3779:
3730:
3684:
3635:
3613:
3578:
3535:
3478:
3432:
3389:Molecular Cell
3375:
3346:(2): a000646.
3326:
3289:(21): R825-8.
3266:
3219:
3180:
3131:
3102:(6): 1155–68.
3082:
3067:
3061:(56): 177–92.
3045:
2995:
2946:
2927:(1–2): 42–66.
2911:
2904:
2881:
2829:
2808:
2790:
2761:(5): 1027–38.
2741:
2734:
2711:
2675:
2624:
2581:
2532:
2479:
2427:
2394:Human Genetics
2377:
2334:
2293:
2286:
2238:
2237:
2235:
2232:
2231:
2230:
2225:
2220:
2215:
2208:
2205:
2132:Rudolf Virchow
2069:salivary gland
2043:
2040:
1924:
1921:
1872:, also called
1828:Multinucleated
1821:Main article:
1818:
1815:
1783:erythropoiesis
1758:
1755:
1742:
1739:
1731:autoantibodies
1722:
1719:
1647:closed mitosis
1595:An image of a
1588:
1585:
1520:Macromolecules
1509:Main article:
1506:
1503:
1501:
1498:
1441:Main article:
1438:
1435:
1427:topoisomerases
1388:Main article:
1385:
1382:
1369:Main article:
1366:
1363:
1347:signal pathway
1305:
1302:
1273:
1270:
1265:Osmotic stress
1240:
1237:
1228:
1225:
1188:Main article:
1185:
1182:
1174:
1171:
1161:
1158:
1132:
1131:
1129:
1126:
1122:
1121:
1119:
1116:
1112:
1111:
1109:
1106:
1102:
1101:
1099:
1096:
1092:
1091:
1089:
1086:
1082:
1081:
1079:
1076:
1072:
1071:
1064:
1059:
1057:Structure name
1044:Nuclear bodies
1042:Main article:
1039:
1036:
991:
988:
950:
947:
908:messenger RNAs
854:tandem repeats
850:nuclear bodies
825:Nuclear bodies
817:Main article:
814:
811:
714:nuclear lamina
704:Nuclear lamina
702:Main article:
699:
698:Nuclear lamina
696:
649:nuclear basket
613:Purkinje cells
539:
536:
512:nuclear matrix
490:(complexed as
461:
458:
338:Main article:
335:
332:
320:nuclear bodies
246:nuclear matrix
175:
174:
172:
171:
166:
161:
156:
142:
137:
132:
127:
122:
116:
111:
106:
100:
95:
88:
80:
79:
71:
70:
43:with the blue
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
7941:
7930:
7927:
7925:
7922:
7920:
7917:
7916:
7914:
7901:
7900:
7893:
7883:
7882:
7877:
7876:
7873:
7867:
7864:
7863:
7860:
7853:
7850:
7847:
7844:
7843:
7840:
7832:
7829:
7827:
7824:
7823:
7822:
7821:
7819:
7816:
7812:
7804:
7801:
7799:
7796:
7795:
7794:
7791:
7787:
7784:
7782:
7779:
7777:
7774:
7772:
7769:
7767:
7764:
7762:
7759:
7757:
7754:
7752:
7749:
7748:
7747:
7744:
7740:
7737:
7735:
7732:
7730:
7727:
7726:
7725:
7722:
7721:
7719:
7716:
7712:
7706:
7703:
7701:
7698:
7696:
7693:
7692:
7689:
7686:
7682:
7676:
7673:
7671:
7668:
7664:
7661:
7659:
7656:
7655:
7654:
7651:
7647:
7644:
7642:
7639:
7637:
7634:
7632:
7629:
7627:
7624:
7622:
7619:
7617:
7614:
7612:
7609:
7608:
7607:
7604:
7603:
7601:
7599:
7595:
7589:
7586:
7582:
7579:
7577:
7574:
7572:
7569:
7567:
7564:
7562:
7559:
7557:
7554:
7552:
7549:
7547:
7544:
7542:
7539:
7537:
7534:
7532:
7529:
7527:
7524:
7522:
7519:
7517:
7514:
7512:
7509:
7507:
7504:
7502:
7499:
7497:
7494:
7492:
7489:
7488:
7487:
7484:
7481:
7478:
7477:
7475:
7473:
7468:
7464:
7460:
7456:
7448:
7443:
7441:
7436:
7434:
7429:
7428:
7425:
7413:
7410:
7408:
7405:
7404:
7402:
7398:
7392:
7389:
7387:
7384:
7380:
7377:
7375:
7372:
7371:
7370:
7367:
7363:
7360:
7358:
7355:
7353:
7350:
7349:
7348:
7345:
7343:
7340:
7339:
7337:
7333:
7327:
7324:
7320:
7317:
7315:
7312:
7310:
7309:Proteinoplast
7307:
7305:
7302:
7300:
7297:
7295:
7292:
7290:
7287:
7285:
7282:
7280:
7277:
7276:
7275:
7272:
7270:
7269:Mitochondrion
7267:
7266:
7264:
7262:
7261:Endosymbionts
7258:
7250:
7247:
7245:
7244:Lamellipodium
7242:
7241:
7240:
7237:
7233:
7230:
7228:
7225:
7223:
7220:
7218:
7215:
7214:
7213:
7210:
7208:
7205:
7201:
7198:
7196:
7193:
7191:
7188:
7186:
7183:
7182:
7181:
7178:
7176:
7173:
7171:
7168:
7166:
7163:
7161:
7160:Microfilament
7158:
7157:
7155:
7153:
7149:
7141:
7138:
7136:
7133:
7131:
7128:
7126:
7123:
7121:
7118:
7117:
7116:
7113:
7109:
7106:
7104:
7101:
7099:
7096:
7094:
7091:
7089:
7086:
7084:
7081:
7080:
7079:
7076:
7074:
7073:Autophagosome
7071:
7069:
7066:
7064:
7061:
7059:
7056:
7054:
7051:
7049:
7048:Cell membrane
7046:
7045:
7043:
7041:
7038:Endomembrane
7035:
7031:
7027:
7019:
7014:
7012:
7007:
7005:
7000:
6999:
6996:
6987:
6983:
6979:
6975:
6973:
6969:
6965:
6962:
6956:
6952:
6948:
6943:
6938:
6934:
6930:
6926:
6921:
6916:
6911:
6907:
6903:
6899:
6895:
6894:"The Nucleus"
6891:
6890:
6884:
6881:
6879:
6876:
6874:
6871:
6870:
6865:
6860:
6851:
6845:
6844:
6839:
6833:
6829:
6824:
6823:
6819:
6818:
6813:
6809:
6805:
6801:
6797:
6793:
6789:
6785:
6780:
6779:
6775:
6774:
6769:
6765:
6761:
6757:
6752:
6747:
6743:
6739:
6732:
6727:
6726:
6722:
6721:
6716:
6712:
6708:
6704:
6700:
6696:
6691:
6690:
6686:
6685:
6680:
6676:
6671:
6666:
6663:(5): 533–47.
6662:
6658:
6654:
6649:
6648:
6643:
6636:
6630:
6624:
6620:
6613:
6611:
6607:
6602:
6595:
6592:
6587:
6581:
6576:
6575:
6566:
6563:
6558:
6554:
6550:
6546:
6542:
6538:
6531:
6528:
6523:
6517:
6513:
6509:
6501:
6496:
6489:
6486:
6481:
6477:
6473:
6469:
6465:
6461:
6458:(4): 513–23.
6457:
6453:
6446:
6443:
6438:
6434:
6430:
6426:
6422:
6418:
6414:
6410:
6403:
6400:
6395:
6391:
6386:
6381:
6377:
6373:
6369:
6365:
6361:
6354:
6351:
6346:
6342:
6338:
6334:
6330:
6326:
6322:
6318:
6315:(5): 419–25.
6314:
6310:
6303:
6300:
6295:
6291:
6287:
6283:
6278:
6273:
6269:
6265:
6261:
6257:
6253:
6246:
6243:
6238:
6234:
6229:
6224:
6219:
6214:
6210:
6206:
6203:(3): 1420–5.
6202:
6198:
6194:
6187:
6184:
6179:
6175:
6171:
6167:
6163:
6159:
6152:
6149:
6144:
6140:
6136:
6132:
6129:(5): 525–33.
6128:
6124:
6117:
6114:
6111:
6105:
6102:
6097:
6093:
6089:
6085:
6081:
6077:
6070:
6067:
6062:
6056:
6052:
6048:
6047:
6042:
6036:
6033:
6028:
6024:
6020:
6013:
6010:
6005:
6001:
5996:
5991:
5987:
5983:
5979:
5975:
5971:
5964:
5961:
5956:
5952:
5947:
5942:
5938:
5934:
5931:(100): 8–15.
5930:
5926:
5922:
5915:
5912:
5907:
5903:
5899:
5895:
5891:
5887:
5883:
5879:
5875:
5869:
5866:
5861:
5857:
5852:
5847:
5842:
5837:
5833:
5829:
5826:(8): e43763.
5825:
5821:
5817:
5810:
5807:
5802:
5798:
5793:
5788:
5784:
5780:
5777:(2): 483–91.
5776:
5772:
5768:
5761:
5758:
5753:
5749:
5744:
5739:
5735:
5731:
5727:
5723:
5719:
5712:
5709:
5704:
5700:
5695:
5690:
5685:
5680:
5676:
5672:
5671:PLOS Genetics
5668:
5661:
5658:
5653:
5649:
5644:
5639:
5635:
5631:
5628:(4): 706–32.
5627:
5623:
5619:
5612:
5609:
5604:
5600:
5596:
5592:
5588:
5584:
5577:
5574:
5569:
5565:
5560:
5555:
5550:
5545:
5541:
5537:
5533:
5529:
5525:
5518:
5515:
5510:
5506:
5501:
5496:
5492:
5488:
5484:
5477:
5474:
5469:
5465:
5461:
5457:
5453:
5449:
5446:(3): 353–62.
5445:
5441:
5434:
5431:
5426:
5422:
5418:
5414:
5411:(1–2): 91–9.
5410:
5406:
5399:
5396:
5391:
5387:
5383:
5379:
5376:(3): 427–36.
5375:
5371:
5364:
5361:
5356:
5350:
5346:
5342:
5335:
5332:
5327:
5323:
5319:
5315:
5311:
5307:
5303:
5299:
5292:
5289:
5284:
5280:
5275:
5270:
5266:
5262:
5258:
5254:
5250:
5243:
5240:
5235:
5231:
5226:
5221:
5217:
5213:
5209:
5205:
5201:
5194:
5191:
5186:
5182:
5178:
5174:
5167:
5165:
5161:
5156:
5152:
5148:
5144:
5139:
5134:
5130:
5126:
5122:
5118:
5114:
5107:
5104:
5099:
5095:
5090:
5085:
5080:
5075:
5071:
5067:
5063:
5056:
5053:
5048:
5042:
5038:
5031:
5028:
5023:
5019:
5015:
5011:
5007:
5003:
4999:
4995:
4988:
4981:
4978:
4973:
4967:
4963:
4956:
4953:
4948:
4942:
4939:. Wiley-VCH.
4938:
4931:
4928:
4923:
4919:
4915:
4911:
4907:
4903:
4896:
4893:
4888:
4884:
4880:
4876:
4873:(1): 607–60.
4872:
4868:
4861:
4858:
4853:
4849:
4845:
4841:
4837:
4833:
4829:
4825:
4818:
4815:
4810:
4804:
4799:
4798:
4789:
4786:
4781:
4777:
4772:
4767:
4762:
4757:
4753:
4749:
4745:
4738:
4735:
4730:
4726:
4721:
4716:
4712:
4708:
4704:
4700:
4696:
4689:
4687:
4683:
4678:
4674:
4669:
4664:
4660:
4656:
4652:
4648:
4644:
4637:
4634:
4629:
4625:
4620:
4615:
4611:
4607:
4603:
4599:
4595:
4588:
4585:
4580:
4576:
4571:
4566:
4562:
4558:
4554:
4550:
4546:
4539:
4536:
4531:
4527:
4522:
4517:
4513:
4509:
4505:
4501:
4497:
4490:
4488:
4484:
4471:
4467:
4460:
4458:
4454:
4449:
4445:
4440:
4435:
4431:
4427:
4423:
4419:
4415:
4408:
4406:
4402:
4397:
4393:
4388:
4383:
4379:
4375:
4371:
4367:
4363:
4356:
4353:
4348:
4344:
4340:
4336:
4332:
4328:
4321:
4318:
4313:
4309:
4304:
4299:
4295:
4291:
4287:
4283:
4279:
4272:
4269:
4264:
4260:
4255:
4250:
4246:
4242:
4239:(4): 919–31.
4238:
4234:
4230:
4223:
4220:
4215:
4211:
4207:
4203:
4199:
4195:
4191:
4187:
4180:
4177:
4172:
4168:
4163:
4158:
4154:
4150:
4147:(5): 637–44.
4146:
4142:
4138:
4131:
4128:
4123:
4119:
4114:
4109:
4105:
4101:
4097:
4093:
4089:
4082:
4080:
4078:
4074:
4069:
4065:
4060:
4055:
4051:
4047:
4042:
4037:
4033:
4029:
4025:
4018:
4016:
4014:
4012:
4008:
4003:
3999:
3994:
3989:
3985:
3981:
3977:
3973:
3969:
3962:
3959:
3954:
3950:
3946:
3942:
3938:
3934:
3931:(2): 181–92.
3930:
3926:
3919:
3916:
3911:
3907:
3902:
3897:
3893:
3889:
3886:(2): 317–21.
3885:
3881:
3877:
3870:
3868:
3866:
3862:
3857:
3851:
3846:
3845:
3836:
3834:
3832:
3828:
3823:
3819:
3815:
3811:
3807:
3803:
3799:
3795:
3788:
3786:
3784:
3780:
3775:
3771:
3766:
3761:
3757:
3753:
3749:
3745:
3741:
3734:
3731:
3726:
3722:
3718:
3714:
3710:
3706:
3703:(3): 209–20.
3702:
3698:
3691:
3689:
3685:
3680:
3676:
3671:
3666:
3662:
3658:
3654:
3650:
3646:
3639:
3636:
3623:
3617:
3614:
3609:
3605:
3601:
3597:
3593:
3589:
3582:
3579:
3574:
3570:
3566:
3562:
3558:
3554:
3551:(3): 345–54.
3550:
3546:
3539:
3536:
3528:
3524:
3520:
3516:
3512:
3508:
3504:
3501:(3): 345–54.
3500:
3496:
3489:
3482:
3479:
3474:
3470:
3466:
3462:
3458:
3454:
3451:(8): 605–12.
3450:
3446:
3439:
3437:
3433:
3428:
3424:
3420:
3416:
3411:
3406:
3402:
3398:
3394:
3390:
3386:
3379:
3376:
3371:
3367:
3362:
3357:
3353:
3349:
3345:
3341:
3337:
3330:
3327:
3322:
3318:
3314:
3310:
3305:
3300:
3296:
3292:
3288:
3284:
3280:
3273:
3271:
3267:
3262:
3258:
3254:
3250:
3246:
3242:
3238:
3234:
3230:
3223:
3220:
3215:
3211:
3207:
3203:
3199:
3195:
3191:
3184:
3181:
3176:
3172:
3167:
3162:
3158:
3154:
3150:
3146:
3142:
3135:
3132:
3127:
3123:
3118:
3113:
3109:
3105:
3101:
3097:
3093:
3086:
3083:
3078:
3074:
3070:
3068:9781134279838
3064:
3060:
3056:
3049:
3046:
3041:
3037:
3032:
3027:
3024:(5): 533–47.
3023:
3019:
3015:
3008:
3006:
3004:
3002:
3000:
2996:
2991:
2987:
2982:
2977:
2973:
2969:
2966:(4): 725–35.
2965:
2961:
2957:
2950:
2947:
2942:
2938:
2934:
2930:
2926:
2922:
2915:
2912:
2907:
2901:
2897:
2890:
2888:
2886:
2882:
2877:
2873:
2869:
2865:
2860:
2855:
2852:(3): 187–98.
2851:
2847:
2843:
2836:
2834:
2830:
2825:
2819:
2811:
2809:9780393680393
2805:
2801:
2794:
2791:
2786:
2782:
2777:
2772:
2768:
2764:
2760:
2756:
2752:
2745:
2742:
2737:
2731:
2727:
2720:
2718:
2716:
2712:
2707:
2700:
2698:
2696:
2694:
2692:
2690:
2688:
2686:
2684:
2682:
2680:
2676:
2671:
2667:
2662:
2657:
2652:
2647:
2643:
2639:
2635:
2628:
2625:
2620:
2616:
2612:
2608:
2604:
2600:
2596:
2592:
2585:
2582:
2577:
2573:
2568:
2563:
2559:
2555:
2551:
2547:
2543:
2536:
2533:
2528:
2524:
2520:
2515:
2510:
2506:
2502:
2498:
2494:
2490:
2483:
2480:
2475:
2471:
2467:
2463:
2458:
2453:
2449:
2445:
2438:
2431:
2428:
2423:
2419:
2415:
2411:
2407:
2403:
2399:
2395:
2388:
2381:
2378:
2373:
2369:
2365:
2361:
2357:
2353:
2349:
2345:
2338:
2335:
2330:
2326:
2321:
2316:
2312:
2308:
2304:
2297:
2294:
2289:
2283:
2278:
2277:
2268:
2266:
2264:
2262:
2260:
2258:
2256:
2254:
2252:
2250:
2248:
2246:
2244:
2240:
2233:
2229:
2226:
2224:
2221:
2219:
2216:
2214:
2211:
2210:
2206:
2204:
2202:
2198:
2194:
2190:
2186:
2182:
2178:
2174:
2170:
2166:
2165:Ernst Haeckel
2162:
2158:
2154:
2150:
2149:fertilization
2146:
2145:Oscar Hertwig
2141:
2138:
2133:
2129:
2124:
2120:
2116:
2111:
2109:
2105:
2101:
2098:
2094:
2089:
2087:
2078:
2074:
2070:
2067:
2066:
2062:Drawing of a
2060:
2053:
2048:
2041:
2039:
2037:
2032:
2030:
2026:
2022:
2018:
2014:
2010:
2006:
2005:
1999:
1997:
1993:
1989:
1984:
1982:
1978:
1974:
1970:
1966:
1962:
1958:
1954:
1950:
1945:
1941:
1937:
1932:
1930:
1922:
1920:
1918:
1914:
1909:
1907:
1903:
1899:
1895:
1891:
1887:
1883:
1879:
1875:
1871:
1868:. In humans,
1867:
1863:
1859:
1855:
1854:
1850:in the genus
1849:
1845:
1841:
1837:
1833:
1829:
1824:
1823:Multinucleate
1816:
1814:
1812:
1808:
1803:
1800:
1796:
1792:
1788:
1784:
1780:
1776:
1772:
1763:
1756:
1754:
1752:
1748:
1740:
1738:
1736:
1732:
1728:
1720:
1718:
1716:
1710:
1707:
1703:
1699:
1693:
1691:
1687:
1683:
1678:
1676:
1672:
1668:
1664:
1660:
1654:
1652:
1648:
1644:
1640:
1636:
1632:
1631:cell division
1624:
1620:
1616:
1612:
1609:
1605:
1601:
1598:
1593:
1586:
1584:
1580:
1576:
1574:
1570:
1566:
1562:
1558:
1554:
1550:
1541:
1537:
1533:
1529:
1525:
1521:
1517:
1512:
1504:
1499:
1497:
1495:
1491:
1486:
1482:
1477:
1475:
1471:
1467:
1463:
1459:
1454:
1450:
1444:
1436:
1434:
1432:
1428:
1424:
1420:
1415:
1407:
1402:
1397:
1391:
1383:
1381:
1379:
1372:
1364:
1362:
1358:
1356:
1352:
1348:
1344:
1340:
1334:
1332:
1328:
1324:
1320:
1316:
1311:
1303:
1301:
1299:
1295:
1291:
1287:
1283:
1279:
1271:
1269:
1266:
1262:
1258:
1254:
1250:
1247:, broken and
1246:
1238:
1236:
1234:
1226:
1224:
1222:
1221:transcription
1218:
1214:
1210:
1206:
1202:
1196:
1191:
1183:
1181:
1179:
1172:
1170:
1168:
1159:
1157:
1155:
1151:
1146:
1144:
1139:
1130:
1128:20–25 nm
1127:
1124:
1123:
1120:
1117:
1114:
1113:
1110:
1107:
1104:
1103:
1100:
1097:
1094:
1093:
1090:
1087:
1084:
1083:
1080:
1077:
1074:
1073:
1063:
1058:
1051:
1045:
1037:
1035:
1033:
1029:
1025:
1020:
1018:
1015:(snoRNA) and
1014:
1010:
1005:
996:
989:
987:
984:
983:
978:
977:B snurposomes
974:
969:
963:
961:
957:
948:
946:
944:
943:nucleophosmin
940:
936:
932:
928:
924:
920:
915:
913:
912:nuclear pores
909:
905:
901:
897:
894:
890:
886:
882:
878:
873:
871:
867:
863:
862:ribosomal RNA
859:
855:
851:
847:
840:
836:
831:
826:
820:
812:
810:
808:
804:
800:
796:
792:
791:
790:laminopathies
785:
783:
779:
778:protofilament
775:
771:
767:
763:
759:
758:alpha-helical
755:
750:
748:
744:
740:
736:
732:
728:
723:
718:
715:
711:
705:
697:
695:
693:
689:
685:
681:
677:
673:
669:
665:
661:
657:
652:
650:
646:
645:nucleic acids
642:
638:
634:
630:
626:
622:
618:
614:
610:
606:
601:
599:
595:
591:
587:
586:nuclear pores
583:
579:
575:
571:
563:
559:
554:
549:
545:
537:
535:
533:
529:
525:
521:
517:
513:
509:
505:
497:
493:
489:
485:
484:nuclear pores
481:
477:
472:
467:
459:
457:
455:
451:
447:
443:
439:
434:
432:
431:
426:
422:
418:
416:
411:
409:
404:
400:
396:
391:
389:
385:
381:
377:
373:
365:
361:
357:
352:
347:
341:
333:
331:
329:
325:
321:
317:
313:
309:
305:
301:
297:
296:nuclear pores
293:
288:
286:
282:
278:
274:
270:
266:
262:
258:
254:
249:
247:
243:
239:
235:
231:
227:
223:
219:
216:
212:
208:
200:
199:
198:
193:
192:
186:
182:
170:
169:Cell membrane
167:
165:
162:
160:
157:
154:
150:
146:
143:
141:
138:
136:
135:Mitochondrion
133:
131:
128:
126:
123:
120:
117:
115:
112:
110:
107:
104:
101:
99:
96:
94:
91:
90:
86:
81:
78:
72:
69:
63:
57:
53:
49:
46:
42:
38:
34:
30:
19:
7924:Cell anatomy
7919:Cell nucleus
7896:
7878:
7480:Pore complex
7455:cell nucleus
7454:
7289:Gerontoplast
7239:Pseudopodium
7232:Radial spoke
7212:Undulipodium
7152:Cytoskeleton
7068:Parenthesome
7052:
6986:the original
6981:
6955:the original
6950:
6937:the original
6932:
6918:Bickmore W.
6909:
6897:
6873:Online books
6864:Cell nucleus
6863:
6828:Cell Biology
6827:
6787:
6783:
6741:
6737:
6698:
6694:
6660:
6656:
6618:
6600:
6594:
6573:
6565:
6540:
6536:
6530:
6507:
6498:
6494:
6488:
6455:
6451:
6445:
6415:(1): 54–63.
6412:
6408:
6402:
6367:
6363:
6353:
6312:
6308:
6302:
6262:(3): 251–6.
6259:
6255:
6245:
6200:
6196:
6186:
6161:
6157:
6151:
6126:
6122:
6116:
6104:
6082:(6): 630–7.
6079:
6075:
6069:
6045:
6035:
6027:the original
6022:
6012:
5977:
5973:
5963:
5928:
5924:
5914:
5881:
5877:
5868:
5823:
5819:
5809:
5774:
5770:
5760:
5725:
5721:
5711:
5674:
5670:
5660:
5625:
5621:
5611:
5589:(2): 233–8.
5586:
5582:
5576:
5531:
5527:
5517:
5490:
5486:
5476:
5443:
5439:
5433:
5408:
5404:
5398:
5373:
5369:
5363:
5344:
5334:
5301:
5297:
5291:
5259:(3): 621–6.
5256:
5252:
5242:
5207:
5203:
5193:
5176:
5172:
5120:
5116:
5106:
5069:
5065:
5055:
5036:
5030:
4997:
4993:
4980:
4964:. Springer.
4961:
4955:
4936:
4930:
4908:(2): 48–52.
4905:
4901:
4895:
4870:
4866:
4860:
4827:
4823:
4817:
4796:
4788:
4751:
4747:
4737:
4702:
4698:
4650:
4646:
4636:
4601:
4597:
4587:
4552:
4549:Open Biology
4548:
4538:
4503:
4499:
4474:. Retrieved
4470:the original
4421:
4417:
4369:
4365:
4355:
4330:
4326:
4320:
4285:
4281:
4271:
4236:
4232:
4222:
4189:
4185:
4179:
4144:
4140:
4130:
4095:
4091:
4031:
4027:
3975:
3971:
3961:
3928:
3924:
3918:
3883:
3879:
3844:Cell Biology
3843:
3797:
3793:
3747:
3743:
3733:
3700:
3696:
3652:
3648:
3638:
3626:. Retrieved
3616:
3594:(1): 19–26.
3591:
3587:
3581:
3548:
3544:
3538:
3498:
3494:
3481:
3448:
3444:
3392:
3388:
3378:
3343:
3339:
3329:
3286:
3282:
3236:
3232:
3222:
3200:(3): 322–7.
3197:
3193:
3183:
3151:(4): 603–8.
3148:
3144:
3134:
3099:
3095:
3085:
3058:
3048:
3021:
3017:
2963:
2959:
2949:
2924:
2920:
2914:
2895:
2849:
2845:
2799:
2793:
2758:
2754:
2744:
2725:
2705:
2641:
2637:
2627:
2597:(2): 384–5.
2594:
2590:
2584:
2549:
2545:
2535:
2527:the original
2496:
2492:
2482:
2447:
2443:
2430:
2400:(4): 281–7.
2397:
2393:
2380:
2350:(1): 53–69.
2347:
2343:
2337:
2310:
2306:
2296:
2275:
2192:
2142:
2128:Robert Remak
2118:
2112:
2100:Robert Brown
2090:
2082:
2063:
2035:
2033:
2013:phagocytosis
2002:
2000:
1996:phagocytosed
1985:
1961:mitochondria
1957:chloroplasts
1953:myxobacteria
1949:methanogenic
1933:
1929:evolutionary
1926:
1911:A number of
1910:
1866:micronucleus
1862:macronucleus
1851:
1826:
1804:
1795:reticulocyte
1791:erythroblast
1768:
1744:
1724:
1711:
1694:
1680:However, in
1679:
1671:prometaphase
1663:microtubules
1655:
1651:open mitosis
1628:
1581:
1577:
1545:
1478:
1466:RNA splicing
1446:
1431:supercoiling
1416:
1412:
1377:
1374:
1359:
1343:inflammatory
1335:
1307:
1275:
1248:
1244:
1242:
1235:processing.
1230:
1197:
1193:
1184:Paraspeckles
1176:
1163:
1147:
1143:paraspeckles
1135:
1115:Paraspeckles
1085:Clastosomes
1075:Cajal bodies
1061:
1056:
1049:
1021:
1004:Cajal bodies
1001:
980:
964:
952:
938:
926:
922:
916:
874:
843:
788:
786:
781:
777:
770:antiparallel
756:contains an
751:
734:
719:
707:
656:karyopherins
653:
648:
625:nucleoporins
602:
567:
558:nuclear pore
548:Nuclear pore
501:
435:
428:
415:constitutive
414:
407:
392:
388:mitochondria
369:
289:
250:
206:
188:
181:cell nucleus
180:
178:
125:Cytoskeleton
97:
67:Cell biology
39:stained for
29:
7854:(Nucleosol)
7852:Nucleoplasm
7715:SMC protein
7705:Paraspeckle
7486:Nucleoporin
7391:Magnetosome
7357:Spliceosome
7284:Chromoplast
7279:Chloroplast
7170:Microtubule
6537:Protoplasma
6362:. Primary.
6254:. Comment.
6195:. Primary.
6164:: 299–328.
5818:. Primary.
5769:. Primary.
5720:. Primary.
5526:. Primary.
5251:. Primary.
5179:(1): 1–77.
4746:. Primary.
4645:. Primary.
4498:. Primary.
4280:. Primary.
4231:. Primary.
4192:(1): 13–9.
4026:. Primary.
3970:. Primary.
3647:. Primary.
3493:. Primary.
3387:. Primary.
3143:. Primary.
3094:. Primary.
2958:. Primary.
2753:. Primary.
2544:. Primary.
2491:. Primary.
2392:. Primary.
2228:Nucleomorph
2159:enters the
2130:(1852) and
2123:Franz Meyen
2093:Franz Bauer
1904:, known as
1902:macrophages
1886:osteoclasts
1844:mycorrhizae
1834:species of
1832:acantharean
1787:bone marrow
1771:erythrocyte
1715:herpesvirus
1709:apoptosis.
1667:centrosomes
1623:chromosomes
1608:fluorescent
1481:spliceosome
1365:Replication
1290:prokaryotes
1282:translation
1253:proteolytic
1239:Clastosomes
1211:except for
1190:Paraspeckle
931:fibrillarin
803:biochemical
766:coiled coil
672:aldosterone
641:vertebrates
617:nucleoplasm
609:glial cells
532:micrometres
520:nucleoplasm
510:called the
494:), and the
438:nucleosomes
425:centromeres
408:facultative
395:Euchromatin
372:chromosomes
334:Chromosomes
261:chromosomes
257:Nuclear DNA
230:osteoclasts
48:Hoechst dye
45:fluorescent
41:nuclear DNA
7929:Organelles
7913:Categories
7793:DNA repair
7386:Proteasome
7379:Inclusions
7326:Nitroplast
7319:Apicoplast
7304:Elaioplast
7299:Amyloplast
7294:Leucoplast
7249:Filopodium
7195:Basal body
7185:Centrosome
7135:Peroxisome
7130:Glyoxysome
7120:Melanosome
7030:organelles
6701:: 607–60.
6603:: 511–514.
6041:Margulis L
5972:. Review.
5923:. Review.
5620:. Review.
5202:. Review.
5064:. Review.
4992:. Review.
4697:. Review.
4596:. Review.
4547:. Review.
4416:. Review.
4364:. Review.
4139:. Review.
4090:. Review.
3972:Chromosoma
3878:. Review.
3800:: 105–31.
3338:. Review.
3281:. Review.
3231:. Review.
3192:. Review.
3057:. Review.
3016:. Review.
2844:. Review.
2638:PLOS Genet
2442:. Review.
2305:. Review.
2234:References
2177:amphibians
2153:sea urchin
2065:Chironomus
2021:poxviruses
1992:chronocyte
1981:G proteins
1888:a type of
1882:myofibrils
1747:eukaryotic
1690:sporozoans
1522:, such as
1458:5' capping
1404:A generic
1394:See also:
1323:Hexokinase
1315:glycolysis
1263:function.
1261:proteasome
1205:interphase
1201:cell cycle
1105:PML bodies
998:Cajal body
743:interphase
444:, such as
376:cell cycle
356:fibroblast
340:Chromosome
281:structured
271:, such as
244:; and the
215:eukaryotic
183:(from
164:Centrosome
149:organelles
52:interphase
37:HeLa cells
7897:see also
7879:see also
7746:Condensin
7695:Chromatin
7598:Nucleolus
7407:Cell wall
7369:Cytoplasm
7342:Nucleolus
7314:Tannosome
7222:Flagellum
7207:Myofibril
7190:Centriole
7125:Microbody
7098:Phagosome
6746:CiteSeerX
6543:: 23–32.
6504:Cited in
6500:1719-1730
6123:BioEssays
6019:"Archaea"
5874:Pennisi E
5583:Mycologia
4754:(1): 18.
4036:CiteSeerX
3628:30 August
3427:233172170
2818:cite book
2591:Neurology
2452:CiteSeerX
2173:Urschleim
2169:phylogeny
2119:cytoblast
2113:In 1838,
1936:symbiotic
1923:Evolution
1898:monocytes
1890:bone cell
1878:syncytium
1838:and some
1777:from the
1698:proteases
1635:apoptosis
1615:metaphase
1569:hydrolyze
1553:exportins
1549:importins
1419:helicases
1213:telophase
1169:(snRNA).
1098:5 μm
935:nucleolin
846:nucleolus
839:nucleolus
819:Nucleolus
813:Nucleolus
807:phenotype
776:called a
676:signaling
594:ribosomes
574:membranes
528:organelle
496:nucleolus
492:chromatin
478:-studded
421:telomeres
399:expressed
384:karyotype
380:chromatin
328:ribosomes
324:nucleolus
242:cytoplasm
226:no nuclei
213:found in
153:cytoplasm
93:Nucleolus
7400:External
7352:Ribosome
7108:Acrosome
7093:Endosome
7088:Lysosome
6964:Archived
6812:83769250
6804:15297641
6715:10611974
6679:11877373
6557:41287948
6472:16953783
6437:16846615
6394:10888648
6345:21200827
6337:11443345
6294:20542871
6286:11523012
6237:11805300
6178:15910279
6143:16615090
6096:16242992
6043:(1981).
6004:26455774
5955:24508984
5906:83769250
5898:15297641
5860:22916303
5820:PLOS ONE
5703:23935529
5603:16894968
5509:24368503
5468:28973947
5425:10708974
5326:44879431
5318:17364135
5283:11331311
5234:23644379
5147:11882878
5098:26793706
5022:23576288
5014:12626338
4922:14731866
4887:10611974
4852:20647022
4844:15667322
4780:28424053
4729:20610547
4677:17954568
4628:32072080
4579:30355755
4530:16148043
4448:20573717
4396:20452955
4347:15240004
4214:29584217
4171:19720872
4122:11368755
4068:12181345
4002:15164213
3953:44941483
3814:16212489
3774:38720076
3765:11164319
3725:30268055
3717:17318225
3679:10588665
3608:16325406
3565:19005234
3527:Archived
3515:19005234
3465:12923522
3419:33823140
3370:20926517
3321:16865665
3313:14588256
3261:20769260
3253:16328431
3214:15145358
3175:11854306
3126:11121432
3077:15565881
3040:11877373
2868:15702987
2785:10831607
2670:28545058
2619:30482028
2364:16506096
2329:15182349
2223:Nucleoid
2207:See also
2181:molluscs
2097:botanist
2017:predator
1973:histones
1944:bacteria
1874:myocytes
1858:Ciliates
1836:protozoa
1799:mutagens
1702:caspases
1686:ciliates
1643:prophase
1528:proteins
1272:Function
1233:pre-mRNA
1125:Speckles
891:, and a
795:progeria
782:filament
774:tetramer
668:cortisol
666:such as
619:and the
476:ribosome
354:A mouse
273:histones
269:proteins
197:nuculeus
159:Lysosome
103:Ribosome
7724:Cohesin
7374:Cytosol
7274:Plastid
7227:Axoneme
7103:Vacuole
7083:Exosome
7078:Vesicle
7053:Nucleus
6784:Science
6768:9554838
6738:Science
6480:5963228
6417:Bibcode
6317:Bibcode
6264:Bibcode
6205:Bibcode
6051:206–227
5995:4640172
5946:4071446
5878:Science
5851:3423374
5828:Bibcode
5801:3027126
5752:3258008
5743:2188835
5694:3731211
5652:1779932
5568:9326623
5536:Bibcode
5460:7141888
5390:5422968
5274:2190567
5225:3720745
5204:Nucleus
5185:7549180
5155:4431000
5125:Bibcode
5089:4707252
4771:5395811
4720:2926751
4668:2118489
4619:7012776
4570:6223218
4521:1266428
4476:6 March
4439:2890200
4387:2857171
4312:9501098
4303:1170524
4263:1955462
4254:2289954
4206:9132135
4162:2742191
4113:1221839
3993:1592132
3945:9671224
3910:9683623
3901:1377332
3822:8807316
3573:6332495
3523:6332495
3473:6439413
3410:8830378
3361:3039535
3291:Bibcode
3166:2174089
3117:2190592
2990:1429833
2981:2289687
2941:9724605
2846:Traffic
2776:2174828
2661:5436639
2611:7854544
2576:4168731
2523:8947544
2514:2121085
2474:9554838
2444:Science
2422:9261461
2414:2416668
2372:6040822
2108:orchids
2042:History
1977:kinases
1940:archaea
1853:Giardia
1785:in the
1700:called
1613:during
1604:stained
1565:GTPases
1474:adenine
1319:glucose
1245:klastos
1219:Pol II
1209:mitosis
904:introns
754:monomer
731:nesprin
629:daltons
621:cytosol
580:and an
304:channel
224:, have
209:) is a
191:nucleus
145:Cytosol
140:Vacuole
109:Vesicle
98:Nucleus
56:mitosis
7776:NCAPH2
7766:NCAPG2
7756:NCAPD3
7751:NCAPD2
7663:CUGBP1
7636:GEMIN7
7631:GEMIN6
7626:GEMIN5
7621:GEMIN4
7616:GEMIN2
7581:NUP214
7576:NUP210
7571:NUP205
7566:NUP188
7561:NUP160
7556:NUP155
7551:NUP153
7546:NUP133
7541:NUP107
7217:Cilium
7040:system
6898:MBInfo
6861:about
6834:
6810:
6802:
6766:
6748:
6713:
6677:
6625:
6582:
6555:
6518:
6478:
6470:
6435:
6392:
6385:112226
6382:
6343:
6335:
6292:
6284:
6235:
6228:122206
6225:
6176:
6141:
6094:
6057:
6002:
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