CHROMOSOMES undergo dynamic behaviors during mitosis to enable the precise separation of the two replicated sister chromatids. It is vital that the replicated sister chromatids are separated successfully. There are two crucial prerequisites for accurate segregation: (1) cohesion between the replicated chromatids must be maintained until anaphase and (2) compaction of the chromosomes into a manageable form, condensation, must be completed prior to metaphase. These processes require two major protein complexes, the cohesin and condensin complexes. Each of these complexes is founded upon a heterodimer of structural maintenance of chromosomes (SMC) proteins, which are chromosome-associated ATPases (Hirano 1998, 2002). Also within each complex are two or three non-SMC subunits, which attribute specific functions to the SMC holocomplex. Despite a similar structural paradigm, the condensin and cohesin complexes are functionally distinct. Although each complex was originally identified for unique ...
Growth of an organ during development and during adaptation in the adult can be controlled by alterations either in the number or the size of cells. The two mechanisms are fundamentally different and require distinct regulation. Rapamycin is a cell growth inhibitor used to treat a number of clinical indications including graft rejection and cancer. The molecular target of rapamycin is a Ser/Thr kinase, called TOR in yeast or mTOR in mammals. The evolutionarily conserved TOR pathway controls key aspects of cellular growth and metabolism. Among these are protein synthesis, ribosome biogenesis, nutrient transport and processing, autophagy and mitochondrial function. mTOR assembles into two distinct multiprotein complexes, termed mTORC1 and mTORC2. mTORC1 consists of raptor (regulatory associated protein of mTOR), mLST8 (mammalian lethal with SEC13 protein 8) and mTOR, and is sensitive to rapamycin. mTORC2 consists of rictor (rapamycin insensitive companion of mTOR), mSIN1, mLST8 (mammalian stress ...
Transcriptional repression is a general mechanism for regulating transcriptional initiation in organisms ranging from yeast to humans. Accurate initiation of transcription from eukaryotic protein-encoding genes requires the assembly of a large multiprotein complex consisting of RNA polymerase II and general transcription factors such as TFIIA, TFIIB, and TFIID. DR1 is a repressor that interacts with the TATA-binding protein (TBP) of TFIID and prevents the formation of an active transcription complex by precluding the entry of TFIIA and/or TFIIB into the preinitiation complex. The protein encoded by this gene is a corepressor of transcription that interacts with DR1 to enhance DR1-mediated repression. The interaction between this corepressor and DR1 is required for corepressor function and appears to stabilize the TBP-DR1-DNA complex. [provided by RefSeq, Jul 2008 ...
We are witnessing tremendous improvements in our understanding of the organization of existence. towards large multiprotein complexes, in particular in eukaryotes, right now calls for a similarly concerted effort to develop and provide new technologies that are urgently required to create in quality and amount the plethora of multiprotein assemblies that form the complexome, and to regularly study their structure and function in the molecular level. Current attempts towards this objective are summarized and examined with this contribution. a two-step process that yields undamaged protein complexes composed of the tagged bait and any connected partners. This method is particularly useful for detecting stable complexes; more transient complexes are not observed, as they tend to dissociate during purification. Two major proteome-wide studies in using the Faucet method have exposed many previously unfamiliar protein relationships and pathway associations [8, 9]. In one study, Gavin genome which ...
Responsible for the deacetylation of lysine residues on the N-terminal part of the core histones (H2A, H2B, H3 and H4). Histone deacetylation plays an important role in transcriptional regulation, cell cycle progression and developmental events. Histone deacetylases act via the formation of large multiprotein complexes (By similarity).
Responsible for the deacetylation of lysine residues on the N-terminal part of the core histones (H2A, H2B, H3 and H4). Histone deacetylation gives a tag for epigenetic repression and plays an important role in transcriptional regulation, cell cycle progression and developmental events. Histone deacetylases act via the formation of large multiprotein complexes ...
This gene encodes a component of the 26S proteasome. The 26S proteasome is a large multiprotein complex that catalyzes the degradation of ubiquitinated intracellular proteins. The encoded protein is a component of the 19S regulatory cap complex of the 26S proteasome and mediates substrate deubiquitination. A pseudogene of this gene is also located on the long arm of chromosome 2 ...
Protein target information for Condensin complex subunit 1 (zebrafish). Find diseases associated with this biological target and compounds tested against it in bioassay experiments.
View mouse Ncapg2 Chr12:116405355-116463532 with: phenotypes, sequences, polymorphisms, proteins, references, function, expression
TY - JOUR. T1 - Binding Specificity of Multiprotein Signaling Complexes Is Determined by Both Cooperative Interactions and Affinity Preferences. AU - Houtman, Jon C.D.. AU - Higashimoto, Yuichiro. AU - Dimasi, Nazzareno. AU - Cho, Sangwoo. AU - Yamaguchi, Hiroshi. AU - Bowden, Brent. AU - Regan, Carole. AU - Malchiodi, Emilio L.. AU - Mariuzza, Roy. AU - Schuck, Peter. AU - Appella, Ettore. AU - Samelson, Lawrence E.. PY - 2004/4/13. Y1 - 2004/4/13. N2 - The generation of multiprotein complexes at receptors and adapter proteins is crucial for the activation of intracellular signaling pathways. In this study, we used multiple biochemical and biophysical methods to examine the binding properties of several SH2 and SH3 domain-containing signaling proteins as they interact with the adapter protein linker for activation of T-cells (LAT) to form multiprotein complexes. We observed that the binding specificity of these proteins for various LAT tyrosines appears to be constrained both by the affinity of ...
proteopedia link. Many PDB files contain complexes in which a particular protein is interacting with a different protein in what are called multi-protein assemblies or multi-protein complexes. These interactions, if biologically relevant, can be immensely insightful in shedding light on cellular and extracellular processes. ...
The Target of Rapamycin (TOR) is a conserved serine/threonine (ser/thr) kinase that functions in two, distinct, multiprotein complexes called TORC1 and TORC2. Each complex regulates different aspects of eukaryote growth: TORC1 regulates cell volume and/or mass by influencing protein synthesis and tu …
The COP9 signalosome (CSN) is a conserved protein complex, typically composed of eight subunits (designated as CSN1 to CSN8) in higher eukaryotes such as ...
Probable protease subunit of the COP9 signalosome complex (CSN), a complex involved in various cellular and developmental processes such as photomorphogenesis and auxin and jasmonate responses. The CSN complex is an essential regulator of the ubiquitin (Ubl) conjugation pathway by mediating the deneddylation of the cullin subunits of the SCF-type E3 ligase complexes, leading to decrease the Ubl ligase activity of SCF. In the complex, it probably acts as the catalytic center that mediates the cleavage of Nedd8 from cullins. It however has no metalloprotease activity by itself and requires the other subunits of the CSN complex (By similarity). The CSN complex is involved in repression of photomorphogenesis in darkness by regulating the activity of COP1-containing Ubl ligase complexes. The complex is also required for degradation of PSIAA6 by regulating the activity of the Ubl ligase SCF-TIR complex. Not involved in CSNs deneddylation/derubylation activity (PubMed:15486099, PubMed:17307927). Essential for
Condensins are pentameric complexes that were initially described as being involved in the dynamics of chromosomes during mitosis. It has been recently established that two related complexes (Condensin I and Condensin II) contribute to this process. An increasing sum of studies, using different approaches in various organisms, leads to the paradigm that Condensins are required for the correct segregation of replicated chromosomes by cooperating somehow with Topoisomerase II in sister chromatid resolution. Depending on species and/or experimental studies, these complexes also contribute to some aspects of the assembly and compaction of mitotic chromosomes. Recent studies provided evidences that Condensins and related complexes also function in non-mitotic processes such as replication and transcription. Biochemical studies have highlighted mechanistic aspects of Condensin function and initiated a fine functional dissection of core and regulatory subunits. However, the exact contribution of each subunit
We used the tracking data to determine the velocity and processivity of wild-type condensin. A plot of the velocity distributions for data collected in the presence of saturated concentrations of ATP (4 mM; Fig. 1G) was well described by a log-normal distribution, revealing a mean apparent translocation velocity of 63 ± 36 base pairs (bp) s−1 (16 ± 9 nm s−1; means ± SD; n = 491) (Fig. 3E). Upon initial binding, condensin paused for a brief period (τpause = 13.3 ± 1.5 s; mean ± SD) before beginning to move along the DNA, which suggests the existence of a rate-limiting step before condensin becomes active for translocation (Fig. 2C and fig. S5). Each translocating condensin complex remained bound to the DNA for an average total time of 4.7 ± 0.2 min and traveled an average of 10.3 ± 0.4 kb (2.6 ± 0.1 μm; means ± SD) before dissociating (Fig. 3F and fig. S6A). These values provide merely a lower limit of the processivity of condensin, because a considerable fraction (42%) of the ...
WD40 domains are abundant in eukaryotes, and they are essential subunits of large multiprotein complexes, which serve as scaffolds. WD40 proteins participate in various cellular processes, such as histone modification, transcription regulation, and signal transduction. WD40 proteins are regarded as crucial regulators of plant development processes. However, the systematic identification and analysis of WD40 proteins have yet to be reported in wheat. In this study, a total of 743 WD40 proteins were identified in wheat, and they were grouped into 5 clusters and 11 subfamilies. Their gene structures, chromosomal locations, and evolutionary relationships were analyzed. Among them, 39 and 46 pairs of TaWD40s were distinguished as tandem duplication and segmental duplication genes. The 123 OsWD40s were identified to exhibit synteny with TaWD40s. TaWD40s showed the specific characteristics at the reproductive developmental stage, and numerous TaWD40s were involved in responses to stresses, including cold, heat
DELLA protein RGA; Probable transcriptional regulator that acts as a repressor of the gibberellin (GA) signaling pathway. Probably acts by participating in large multiprotein complexes that repress transcription of GA-inducible genes. Positively regulates XERICO expression in seeds. Upon GA application, it is degraded by the proteasome, allowing the GA signaling pathway. Compared to other DELLA proteins, it is the most sensitive to GA application. No effect of the BOI proteins on its stability. Its activity is probably regulated by other phytohormones such as auxin and ethylene, attenu [...] (587 aa ...
The mechanisms by which nuclear receptors transmit a hormone binding signal to core transcription machinery have been the focus of intensive research. These efforts have lead to the identification and characterization of several distinct multiprotein complexes which directly interact with agonist-bound nuclear receptors (9, 16, 28, 45, 46). The SRC-1 family of nuclear receptor coactivators appear to play a critical role in mediating the association of one of these complexes to nuclear receptors. This complex has been shown to contain potent HAT activities in p300-CBP and also the p300/CBP-associated factor PCAF. Additionally, these factors have been identified in complexes that contain intrinsic chromatin remodeling activity. These findings suggest that the SRC-1-containing complex may have as its primary purpose the chemical modification of the chromatin surrounding a target gene to which it is recruited. A second complex, identified on the basis of its ability to interact with activated ...
InterPro provides functional analysis of proteins by classifying them into families and predicting domains and important sites. We combine protein signatures from a number of member databases into a single searchable resource, capitalising on their individual strengths to produce a powerful integrated database and diagnostic tool.
Background Component of the COP9 signalosome complex (CSN), a complex involved in various cellular and developmental processes. The CSN complex is an essential regulator of the ubiquitin (Ubl) conjugation pathway by mediating...
Losada, A., Yokochi, T., Hirano, T. (May 2005) Functional contribution of Pds5 to cohesin-mediated cohesion in human cells and Xenopus egg extracts. J Cell Sci, 118 (Pt 10). pp. 2133-41. ISSN 0021-9533 (Print) Ono, T., Losada, A., Hirano, M., Myers, M. P., Neuwald, A. F., Hirano, T. (October 2003) Differential contributions of condensin I and condensin II to mitotic chromosome architecture in vertebrate cells. Cell, 115 (1). pp. 109-121. ISSN 0092-8674 Losada, A., Hirano, M., Hirano, T. (December 2002) Cohesin release is required for sister chromatid resolution, but not for condensin-mediated compaction, at the onset of mitosis. Genes & Development, 16 (23). pp. 3004-3016. ISSN 0890-9369 MacCallum, D. E., Losada, A., Kobayashi, R., Hirano, T. (January 2002) ISWI remodeling complexes in Xenopus egg extracts: Identification as major chromosomal components that are regulated by INCENP-aurora B. Molecular Biology of the Cell, 13 (1). pp. 25-39. ISSN 1059-1524 Losada, A., Hirano, T. (October 2001) ...
InterPro provides functional analysis of proteins by classifying them into families and predicting domains and important sites. We combine protein signatures from a number of member databases into a single searchable resource, capitalising on their individual strengths to produce a powerful integrated database and diagnostic tool.
This gene encodes a subunit of the condensin complex, which is responsible for the condensation and stabilization of chromosomes during mitosis and meiosis. Phosphorylation of the encoded protein activates the condensin complex. There are pseudogenes for this gene on chromosomes 8 and 15. Alternative splicing results in multiple transcript variants. [provided by RefSeq, Aug 2012] ...
Mammalian cell growth is tightly linked to adequate supplies of growth factors and nutrients, including glucose and amino acids. The mechanistic target of rapamycin complex 1 (mTORC1) functions as a coincidence detector that supports anabolic metabolism when convergent growth factor- and nutrient-derived signals trigger mTORC1 kinase activation. Conversely, nutrient starvation suppresses mTORC1 activity and triggers a shift to catabolic pathways, such as autophagy, to support cell survival under austere conditions. An elaborate signaling apparatus has evolved to harmonize mTORC1 kinase activation and protein synthesis with supplies of leucine and other amino acids (1). Earlier findings implicated the leucyl-transfer RNA (tRNA) synthetase 1 (LARS1) as a proximate sensor of leucine availability (2). On page 205 of this issue, Yoon et al. (3) report that glucose modulates the functions of LARS1 in leucine sensing and disposition, thereby coordinating leucine-dependent mTORC1 activation and protein ...
The mechanistic target of rapamycin complex 1 (mTORC1) is a central regulator of cell growth and proliferation that is often aberrantly activated in cancer, as...
Background In addition to serving as building blocks for protein synthesis, amino acids also provide energy and precursors that are used by cells through catabolism. Mechanistic target of rapamycin complex 1 (mTORC1) is a central coordinator of cellular metabolism. However, little is known regar...
概要:哺乳动物雷帕霉素靶蛋白复合物1(mTORC1)能够感受一系列细胞内外的环境因素(如氨基酸),从而控制细胞生长和代谢.在过去的几十年里,众多蛋白被发现能够参与受氨基酸调控的mTORC1信号通路中.Rag GTPases能够将氨基酸的信号传递给mTORC1并招募mTORC1到溶酶体表面.近年来,参与mTORC1信号通路的蛋氨酸代谢物、亮氨酸以及精氨酸的感受体逐渐被发现.感受体的鉴定有助于理解细胞是如何通过调整内部氨基酸感应通路来满足自身需求.本文综述了氨基酸调控mTORC1信号通路的分子机制,并探讨了感受体如何将特定氨基酸信号精确传递给mTORC1信号通路 ...
DEPTOR [DEP-domain-containing and mTOR (mammalian target of rapamycin)-interacting protein] is a modulator of mTOR signalling that binds to mTORC (mTOR complex) 1 and mTORC2. However, to date, the precise functions of DEPTOR are not fully elucidated,
HCAP-G antibody (non-SMC condensin I complex, subunit G) for WB. Anti-HCAP-G pAb (GTX131128) is tested in Human samples. 100% Ab-Assurance.
Multiprotein complexes are an emerging focus in current biology, resulting in a demand for advanced heterologous expression systems
The inflammasome is a multi-protein complex which when activated regulates caspase-1 activation and IL-1β and IL-18 secretion. The NLRP3 (NACHT, LRR, and pyrin ...
Interaction between NBS1 and the mTOR/Rictor/SIN1 complex.A & B. Co-immunoprecipitation assays showed the interaction between NBS1 and mTOR in 293T cells ov
UNFCCC COP21大會期間,編輯室與特約記者、合作媒體ENS現場連線,並與譯者合作盤整談判重點做成即時報導,傳遞第一手消息,文章將同步與台達電子文教基金會低碳生活部落格、CNEX 紀實頻道共同刊登。此為「COP21連線」四項子專題之四。各項子專題請見 COP21翻譯米糕|COP21×社群觀點|COP21會前情勢|COP21大會連線
In a blind test of protein-docking algorithms, six groups used different methods to predict the structure of a protein complex. All six predicted structures were close enough to the experimental complex to be useful; nevertheless, several important details of the experimental complex were missed or …
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Histones are modified at specific positions on their conserved amino‐terminal tails, and these modifications play central roles in gene regulation (Jenuwein and Allis, 2001; Turner, 2002). For example, acetylation at lysine 14 of histone H3 (H3K14Ac) or methylation of lysine 4 (H3K4Me) is associated with gene activation. In contrast, the lack of acetylation as well as the methylation of lysine 9 of histone H3 (H3K9Me) is correlated with gene repression (Jenuwein and Allis, 2001; Turner, 2002). These two types of modifications are performed by histone acetyltransferases (HATs) and histone methyltransferases (HMTases). These modifying enzymes are members of large multiprotein complexes with other subunits likely serving roles in targeting or regulation. The primary substrates for these enzymes are the amino‐terminal tails of the histone proteins. Modified tails function as binding platforms for transcriptional regulatory factors. For example, the histone H3 tail methylated at lysine 9 is ...
Chloroplasts originated from an endosymbiotic event in which a free-living cyanobacterium was engulfed by an ancestral eukaryotic host. During evolution the majority of the chloroplast genetic information was transferred to the host cell nucleus. As a consequence, proteins formerly encoded by the chloroplast genome are now translated in the cytosol and must be subsequently imported into the chloroplast. This process involves three steps: (i) cytosolic sorting procedures, (ii) binding to the designated receptor-equipped target organelle and (iii) the consecutive translocation process. During import, proteins have to overcome the two barriers of the chloroplast envelope, namely the outer envelope membrane (OEM) and the inner envelope membrane (IEM). In the majority of cases, this is facilitated by two distinct multiprotein complexes, located in the OEM and IEM, respectively, designated TOC and TIC. Plants are constantly exposed to fluctuating environmental conditions such as temperature and light ...
The mammalian target of rapamycin complex 1 (mTORC1) is mixed up in cellular transcription and translation processes. in control (P) bodies a niche site for mRNA degradation. Incubation of cells with rapamycin a known inhibitor of mTOR kinase activity improved the full total Edc4 proteins expression but at the same time reduced the Edc4 discussion with mTORC1. Furthermore rapamycin treatment led to a significant reduction in total serine phosphorylated Edc4 proteins signal and the full total 5′-capped mRNA. These results provide the 1st proof for the pivotal part of mTORC1 in Edc4 rules. Further in-depth research must get a full knowledge of molecular crosstalk between mTORC1 signaling and mRNA decapping pathway. and axis. Statistical relationship using Pearsons technique [22] for just two 3rd party tests demonstrated coefficients of 0.863 and 0.754 respectively which recommended a high level of co-occurrence of the raptor element of Edc4 and mTORC1. Likewise the Manders overlap coefficients ...
Two simple models can be envisaged: either cohesins are needed to activate condensin function or, alternatively, cohesins are required to ensure correct chromosome folding by condensins. These models can be distinguished by following the state of the mitotic chromosomes after a loss of cohesin activity. In the first scenario, the chromosomes remain in an interphase state, and thus would condense upon the readdition of cohesin and the subsequent activation of condensin. In contrast, the latter scenario predicts that misfolded chromosomes would result from the inappropriate action of condensin, and these would likely prove refractory to refolding. To test this, we asked whether chromosome condensation is reversible in the cohesin mutant mcd1-1. In contrast to both the brn1-9 and ycg1-2 condensin mutants, the condensation defect in the mcd1-1 strain was not reversible (Fig. 7 B). One trivial explanation is that no new functional Mcd1-1p protein is made after the shift to the permissive ...
The mechanistic (or mammalian) target of rapamycin complex 1 (mTORC1) controls cell growth, proliferation, and metabolism in response to diverse stimuli. Two major parallel pathways are implicated in mTORC1 regulation including a growth factor-responsive pathway mediated via TSC2/Rheb and an amino acid-responsive pathway mediated via the Rag GTPases. Here, we identify and characterize three highly conserved growth factor-responsive phosphorylation sites on RagC, a component of the Rag heterodimer, implicating cross talk between amino acid and growth factor-mediated regulation of mTORC1. We find that RagC phosphorylation is associated with destabilization of mTORC1 and is essential for both growth factor and amino acid-induced mTORC1 activation. Functionally, RagC phosphorylation suppresses starvation-induced autophagy, and genetic studies in reveal that RagC phosphorylation plays an essential role in regulation of cell growth. Finally, we identify mTORC1 as the upstream kinase of RagC on S21. ...
mTORC1 signalling promotes SREBP activation and lipogenesis in response to both physiological and genetic stimuli. In primary rodent hepatocytes and the intact liver, insulin or feeding has been shown to increase the expression of the major liver isoform of SREBP (SREBP1c) and its targets, and to promote de novo lipid synthesis in a manner that is sensitive to rapamycin [17,18,19]. Insulin activates mTORC1 through a pathway involving the Akt‐mediated phosphorylation and inhibition of TSC2, within a complex with TSC1 and TBC1D7 ]2,3,4]. Expression of constitutively active Akt or loss of either TSC1 or TSC2, both of which result in insulin‐independent activation of mTORC1 signalling, stimulates the global expression of SREBP1 and SREBP2 targets and drives lipogenesis through mTORC1 [15,16]. These latter studies found that mTORC1 signalling promotes accumulation of the processed, mature form of SREBP1, which resides in the nucleus to induce its own expression and that of genes involved in both ...
Mass spectrometry has proven to be a valuable tool for analyzing large protein complexes. This method enables insights into the...
The COP9 signalosome is a highly conserved protein complex composed of eight subunits. In this study a novel, regulatory mechanism of CSN biogenesis was identified. We used stable transfected siCSN1 cells in which the protein and the mRNA expression of CSN subuntis were downregulated. Transfection of His-CSN1 in those siCSN1 cells led to the induction of the de novo Synthesis of the whole CSN complex. In addition the expression of the transcription factors STAT1 and c-Myc was elevated. The cells were treated with IFN alpha or IFN gamma, respectively. This resulted in the induction of the CSN de novo synthesis. Moreover, the siRNA-mediated inhibition of STAT1, c-Myc, Lin28B as well as treatment with the pharmacological inhibitors AG9 or AG490 led to a reduced protein expression of the analysed CSN subunits. We found that in all experiments there was a significant change on protein level in contrast to a marginal change on the RNA level. Based on our study we hypothesized that the CSN biogenesis ...
Proline-rich AKT1 substrate 1 40 kDa (PRAS40) is a protein encoded by the AKT1S1 gene in humans. PRAS40 binds to 14-3-3 proteins when phosphorylated. PRAS40 is a subunit of the mammalian target of rapamycin complex 1 (mTORC1), and it negatively regulates mTOR activity in a manner that is dependent on its phosphorylation state and binding to 14-3-3 proteins. It inhibits RHEB-GTP-dependent mTORC1 activation. Although it is a substrate for AKT1 phosphorylation, PRAS40 can also be activated by AKT1-independent mechanisms. PRAS40 is also known as AKT1 substrate 1 (proline-rich), AKT1S1, proline-rich AKT1 substrate 1, lobe, and MGC2865.. ...
Proline-rich AKT1 substrate 1 40 kDa (PRAS40) is a protein encoded by the AKT1S1 gene in humans. PRAS40 binds to 14-3-3 proteins when phosphorylated. PRAS40 is a subunit of the mammalian target of rapamycin complex 1 (mTORC1), and it negatively regulates mTOR activity in a manner that is dependent on its phosphorylation state and binding to 14-3-3 proteins. It inhibits RHEB-GTP-dependent mTORC1 activation. Although it is a substrate for AKT1 phosphorylation, PRAS40 can also be activated by AKT1-independent mechanisms. PRAS40 is also known as AKT1 substrate 1 (proline-rich), AKT1S1, proline-rich AKT1 substrate 1, lobe, and MGC2865.. ...
The COP9 signalosome (CSN) is a conserved multiprotein complex, with an important developmental role in several organisms, ranging from plants to mammalians.
NYU Langone study suggests that placing strict biological controls on a portion of a protein complex could help stop a variety of cancers. Learn more.
part of a multiprotein complex, SRCAP-containing complex supporting ATP-dependent exchange of histone dimers containing H2B and H2AFZ into mononucleosomes reconstituted with recombinant H2A, H2B, H3, and H4 ...
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https://doi.org/10.18632/oncotarget.3044 Long Chen, Baoshan Xu, Lei Liu, Chunxiao Liu, Yan Luo, Xin Chen, Mansoureh Barzegar, Jun Chung, Shile Huang
Nuclear fission, or nucleopore, is a large protein complex that passes through a nucleolem (a double membrane that encloses a nucleus in a e
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