Cell nucleus - Knowledge (XXG)

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.

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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,

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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.

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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

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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

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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.

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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

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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

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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

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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

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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

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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.

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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

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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.

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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

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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

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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.

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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.

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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

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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

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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

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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

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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

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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

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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.

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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.

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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 ("

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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

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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.

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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

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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.

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Bernstein, H., Bernstein, C. (2017). Sexual Communication in Archaea, the Precursor to Eukaryotic Meiosis. In: Witzany, G. (eds) Biocommunication of Archaea. Springer, Cham.

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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

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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

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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

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bacteria that possess a nuclear structure with primitive pores and other compartmentalized membrane structures. A similar proposal states that a eukaryote-like cell, the

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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

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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.

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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

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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

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A university level textbook focusing on cell biology. Contains information on nucleus structure and function, including nuclear transport, and subnuclear domains

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where it interacts with transcription factors to downregulate the production of certain enzymes in the pathway. This regulatory mechanism occurs in the case of

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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

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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

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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.

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A review article about the nucleus, explaining the structure of chromosomes within the organelle, and describing the nucleolus and other subnuclear bodies

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Zimber A, Nguyen QD, Gespach C (October 2004). "Nuclear bodies and compartments: functional roles and cellular signalling in health and disease". Review.

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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

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Grigoryev SA, Bulynko YA, Popova EY (2006). "The end adjusts the means: heterochromatin remodelling during terminal cell differentiation". Review.

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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

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for heredity. The function of the nucleus as carrier of genetic information became clear only later, after mitosis was discovered and the

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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.

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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: 2807: 1452: 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.

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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: 6946: 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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speckles increases the kinetic efficiency of pre-mRNA splicing, ultimately boosting protein levels by modulation of splicing.

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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

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Tripathi K, Parnaik VK (September 2008). "Differential dynamics of splicing factor SC35 during the cell cycle". Primary.

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Barned S, Goodman AD, Mattson DH (February 1995). "Frequency of anti-nuclear antibodies in multiple sclerosis". Primary.

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Horton TR (2006). "The number of nuclei in basidiospores of 63 species of ectomycorrhizal Homobasidiomycetes". Primary.

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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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A review article about nuclear transport, explains the principles of the mechanism, and the various transport pathways

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with discrete domains defined by dense localization of the transcription factor PTF, which promotes transcription of

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is stained blue. The distinct chromosome territories of chromosome 2 (red) and chromosome 9 (green) are stained with

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Website covering structure and function of the nucleus from the Department of Oncology at the University of Alberta.

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Goebel HH, Warlo I (January 1997). "Nemaline myopathy with intranuclear rods--intranuclear rod myopathy". Review.

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in DNA, helping it wind and unwind, as well as a large variety of transcription factors that regulate expression.

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Mutations in lamin genes leading to defects in filament assembly cause a group of rare genetic disorders known as

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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: 6450:

de Roos AD (2006). "The origin of the eukaryotic cell based on conservation of existing interfaces". Primary.

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contains digitized commentaries and links to seminal research papers on the nucleus. Published online in the

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Handwerger KE, Gall JG (January 2006). "Subnuclear organelles: new insights into form and function". Review.

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conditions within the nucleus and degrade once there is a decrease in activity or if cells are treated with

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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.

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Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P, eds. (2002). "Chapter 4: DNA and Chromosomes".

32: 7114: 6745: 4545:"Molecular dissection of nuclear paraspeckles: towards understanding the emerging world of the RNP milieu" 4035: 2451: 2217: 1539: 1023: 709: 581: 577: 507: 479: 429: 382:, and during cell division the chromatin can be seen to form the well-defined chromosomes familiar from a 6963: 7923: 7918: 7437: 7378: 7164: 7057: 6634: 2196: 1638: 1560: 1281: 687: 589: 449: 7411: 6893: 6882: 6511: 6416: 6316: 6263: 6204: 5827: 5535: 5483:"The angiosperm phloem sieve tube system: a role in mediating traits important to modern agriculture" 5171:

Boulikas T (1995). "Phosphorylation of transcription factors and control of the cell cycle". Review.

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the molecule guanosine triphosphate (GTP) to release energy. The key GTPase in nuclear transport is

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bound by exportins. The ability of importins and exportins to transport their cargo is regulated by

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membrane. As part of the endoplasmic reticulum membrane, the outer nuclear membrane is studded with

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Lamond AI, Spector DL (August 2003). "Nuclear speckles: a model for nuclear organelles". Review.

3422: 3316: 3256: 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".

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Lodish H, Berk A, Matsudaira P, Kaiser CA, Krieger M, Scott MP, Zipursky SL, Darnell J (2004).

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in modern poxviruses and eukaryotes. It has been suggested that the unresolved question of the

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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

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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

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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

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Fuerst JA (2005). "Intracellular compartmentation in planctomycetes". Review.

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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

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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".

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Watson JD, Baker TA, Bell SP, Gann A, Levine M, Losick R (2004). "Ch9–10".

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Goldman AE, Moir RD, Montag-Lowy M, Stewart M, Goldman RD (November 1992).

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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).

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Matera AG (August 1998). "Of coiled bodies, gems, and salmon". Review.

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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 speckles 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 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