Basic leucine zipper and W2 domain-containing protein 2: Difference between revisions
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'''Basic leucine zipper and W2 domain-containing protein 2''' is a |
'''Basic leucine zipper and W2 domain-containing protein 2''' is a protein that is encoded by the ''BZW2'' gene.<ref name="pmid11042152">{{cite journal | vauthors = Zhang QH, Ye M, Wu XY, Ren SX, Zhao M, Zhao CJ, Fu G, Shen Y, Fan HY, Lu G, Zhong M, Xu XR, Han ZG, Zhang JW, Tao J, Huang QH, Zhou J, Hu GX, Gu J, Chen SJ, Chen Z | title = Cloning and functional analysis of cDNAs with open reading frames for 300 previously undefined genes expressed in CD34+ hematopoietic stem/progenitor cells | journal = Genome Res | volume = 10 | issue = 10 | pages = 1546–60 |date=Nov 2000 | pmid = 11042152 | pmc = 310934 | doi =10.1101/gr.140200 }}</ref><ref name="entrez">{{cite web | title = Entrez Gene: BZW2 basic leucine zipper and W2 domains 2| url = https://www.ncbi.nlm.nih.gov/sites/entrez?Db=gene&Cmd=ShowDetailView&TermToSearch=28969| accessdate = }}</ref> It is a eukaryotic translation factor and is conserved up to [[fungus]], and further conserved in a couple [[bacteria]] species. |
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== Gene == |
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BZW2 is known as Basic Leucine Zipper W2 Domain-Containing Protein 2, MST017, MSTP017, 5MP1, and HSPC028.<ref name=":0">{{Cite web|title=BZW2 basic leucine zipper and W2 domains 2 [Homo sapiens (human)] - Gene - NCBI|url=https://www.ncbi.nlm.nih.gov/gene/28969|access-date=2020-07-31|website=www.ncbi.nlm.nih.gov}}</ref> It is located on chromosome 7 at the 21.1 locus. The gene spans 60,389 base pairs, at coordinates 16,583,248 – 16,804,999. There are 12 exons. |
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== Protein == |
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There are two known [[Protein isoform|isoforms]] of BZW2. Isoform 1 is 419 amino acids long and is the most abundant form. Isoform 2 is 225 amino acids, containing only 11 exons and a shorter [[N-terminus]].<ref name=":0" /> |
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The coded protein is 419 amino acids long and weighs 48.3 kDa.<ref>{{Cite web|title=BZW2 Antibody (PA5-21808)|url=https://www.thermofisher.com/antibody/product/BZW2-Antibody-Polyclonal/PA5-21808|access-date=2020-07-31|website=www.thermofisher.com}}</ref> As described in the name, the protein contains a [[Leucine zipper|leucine-zipper]] motif. Four “L……” repeats are present in the beginning, giving rise to the characteristic leucine zipper helix within the 3D structure. An eIF5C domain follows the leucine motif. |
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BZW2 has a higher amount of [[Lysine|Lysines]] and a lower amount of [[Proline|Prolines]]. There are no significant charge segments.<ref>{{Cite journal|last=Madeira|first=Fábio|last2=Park|first2=Young mi|last3=Lee|first3=Joon|last4=Buso|first4=Nicola|last5=Gur|first5=Tamer|last6=Madhusoodanan|first6=Nandana|last7=Basutkar|first7=Prasad|last8=Tivey|first8=Adrian R N|last9=Potter|first9=Simon C|last10=Finn|first10=Robert D|last11=Lopez|first11=Rodrigo|date=2019-04-12|title=The EMBL-EBI search and sequence analysis tools APIs in 2019|url=http://dx.doi.org/10.1093/nar/gkz268|journal=Nucleic Acids Research|volume=47|issue=W1|pages=W636–W641|doi=10.1093/nar/gkz268|issn=0305-1048}}</ref> |
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The [[Biomolecular structure|secondary structure]] contains a majority of alpha helices. The tertiary structure forms a repeated fold of alpha-helices, a structure that is conserved through bacteria.<ref>{{Cite journal|last=Kelley|first=Lawrence A|last2=Mezulis|first2=Stefans|last3=Yates|first3=Christopher M|last4=Wass|first4=Mark N|last5=Sternberg|first5=Michael J E|date=2015-05-07|title=The Phyre2 web portal for protein modeling, prediction and analysis|url=http://dx.doi.org/10.1038/nprot.2015.053|journal=Nature Protocols|volume=10|issue=6|pages=845–858|doi=10.1038/nprot.2015.053|issn=1754-2189}}</ref> |
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[[File:Predicted Structure of BZW2.jpg|thumb|413x413px|BZW2 structure from Phyre2 colored from N-terminus (red) to C-terminus (blue).]] |
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== Regulation == |
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=== Gene-Level === |
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There are three known [[Promoter|promoters]] for BZW2.<ref>{{Cite web|title=Genomatix - NGS Data Analysis & Personalized Medicine|url=https://www.genomatix.de/|access-date=2020-07-31|website=www.genomatix.de}}</ref> It is regulated by numerous [[Transcription factor|transcription factors]], including an [[Estrogen receptor beta|estrogen receptor]] transcription factor, leucine zipper transcription factor, and Y sex-determining transcription factors. Overall BZW2 is ubiquitously expressed throughout tissues. There is increased [[Messenger RNA|mRNA]] abundance in the [[heart]], [[placenta]], and [[skeletal muscle]]. |
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=== Transcript-Level === |
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There are four major [[Stem-loop|stem loops]] in the 5’ untranslated and four in the 3’ untranslated region.<ref>{{Cite journal|last=Zuker|first=M.|date=2003-07-01|title=Mfold web server for nucleic acid folding and hybridization prediction|url=http://dx.doi.org/10.1093/nar/gkg595|journal=Nucleic Acids Research|volume=31|issue=13|pages=3406–3415|doi=10.1093/nar/gkg595|issn=1362-4962}}</ref> |
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=== Protein-Level === |
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BZW2 is localized in the [[cytoplasm]] and has multiple [[phosphorylation]], [[acetylation]], [[glycosylation]], [[SUMO protein|SUMOylation]] and [[glycation]] sites for regulation.<ref>{{Cite web|title=BZW2 (human)|url=https://www.phosphosite.org/proteinAction.action?id=10754&showAllSites=true|access-date=2020-07-31|website=www.phosphosite.org}}</ref> |
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[[File:BZW2 Post-Translational Modification Schematic.jpg|thumb|646x646px|Schematic of the post-translational modification locations on the BZW2 protein. ]] |
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[[File:BZW2 Sequence Detailing Spacing, Charge Clusters, and Repeats.jpg|thumb|620x620px|Sequence of BZW2 is shown with positions of spacing information, charge clusters, and repeats. ]] |
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== Evolution == |
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BZW2 has a single paralog, [[BZW1]] which is conserved up to plants.<ref>{{Cite web|title=Protein BLAST: search protein databases using a protein query|url=https://blast.ncbi.nlm.nih.gov/Blast.cgi?PROGRAM=blastp&PAGE_TYPE=BlastSearch&LINK_LOC=blasthome|access-date=2020-07-31|website=blast.ncbi.nlm.nih.gov}}</ref> There are BZW2 [[Sequence homology|orthologs]] up to a couple species of bacteria. The most distant ortholog was ''[[Microbacterium|Microbacterium aborescens]]''. BZW2 contains an eIF5C domain which is also present in [[EIF2B2|eIF2BE]], [[EIF4G2|eIF4G]], [[eIF5]], and a GAP protein specific for eIF2.<ref>{{Cite journal|last=Koonin|first=Eugene V.|date=1995-08|title=Multidomain organization of eukaryotic guanine nucleotide exchange translation initiation factor eIF-2B subunits revealed by analysis of conserved sequence motifs|url=http://dx.doi.org/10.1002/pro.5560040819|journal=Protein Science|volume=4|issue=8|pages=1608–1617|doi=10.1002/pro.5560040819|issn=0961-8368}}</ref> |
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Compared to [[Cytochrome c|Cytochrome C]], a quickly diverging protein, and [[Fibronectin]], a slowly diverging protein, BZW2 has had slow corrected divergence over time. |
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[[File:BZW2 Divergence Rate.jpg|thumb|395x395px|Corrected rate of divergence of BZW2 in relation to Cytochrome C and Fibrinogen ]] |
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== Interactions == |
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BZW2 is known to interact with: |
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* [[BZW1]] |
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* [[EIF2S2]] |
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* [[PSTPIP1]] |
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* NEK4 |
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* ORF4 |
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* [[SNW1]] |
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== Clinical Significance == |
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=== Cancer === |
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BZW2 has been studied to determine its role in multiple cancers. Overall, the studies all showed that upregulation of BZW2 lead to more severe forms of cancer, higher rate of mortality, and increased likeliness of reoccurrence. |
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A 2019 study focused on the effect of BZW2 in [[Colorectal cancer|colorectal cancer.]]<ref>{{Cite journal|last=Sato|first=Kuniaki|last2=Masuda|first2=Takaaki|last3=Hu|first3=Qingjiang|last4=Tobo|first4=Taro|last5=Gillaspie|first5=Sarah|last6=Niida|first6=Atsushi|last7=Thornton|first7=Mackenzie|last8=Kuroda|first8=Yousuke|last9=Eguchi|first9=Hidetoshi|last10=Nakagawa|first10=Takashi|last11=Asano|first11=Katsura|date=2019-06|title=Novel oncogene 5MP1 reprograms c-Myc translation initiation to drive malignant phenotypes in colorectal cancer|url=http://dx.doi.org/10.1016/j.ebiom.2019.05.058|journal=EBioMedicine|volume=44|pages=387–402|doi=10.1016/j.ebiom.2019.05.058|issn=2352-3964}}</ref> It found that upregulation of BZW2 promoted [[Neoplasm|tumor]] growth and had a downstream upregulation effect on [[Myc|c-Myc]], a [[Oncogene|proto-oncogene]]. A second study from 2020 determined this upregulation also had a positive effect on the activation of the [[Mitogen-activated protein kinase|ERK/MAPK]] pathway. <ref>{{Cite journal|last=Huang|first=Longping|last2=Chen|first2=Si|last3=Fan|first3=Haijun|last4=Ai|first4=Fan|last5=Sheng|first5=Weiwei|date=2019-10-23|title=BZW2 promotes the malignant progression of colorectal cancer via activating the ERK/MAPK pathway|url=http://dx.doi.org/10.1002/jcp.29361|journal=Journal of Cellular Physiology|volume=235|issue=5|pages=4834–4842|doi=10.1002/jcp.29361|issn=0021-9541}}</ref> |
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In [[hepatocellular carcinoma]], [[Osteosarcoma|ostersarcoma]], [[Adenocarcinoma of the lung|lung adenocarcinoma]], and [[Bladder cancer|muscle-invasive bladder cancer]], overexpression of BZW2 lead to overactivation of the AKT/mTOR signaling pathway by increasing phosphorylation of [[Protein kinase B|AKT]] and [[mTOR]].<ref>{{Cite journal|last=Cheng|first=Dong-Dong|last2=Li|first2=Shi-Jie|last3=Zhu|first3=Bin|last4=Yuan|first4=Ting|last5=Yang|first5=Qing-Cheng|last6=Fan|first6=Cun-Yi|date=2017-04|title=Downregulation of BZW2 inhibits osteosarcoma cell growth by inactivating the Akt/mTOR signaling pathway|url=http://dx.doi.org/10.3892/or.2017.5890|journal=Oncology Reports|volume=38|issue=4|pages=2116–2122|doi=10.3892/or.2017.5890|issn=1021-335X}}</ref><ref>{{Cite journal|last=Jin|first=Xin|last2=Liao|first2=Mingmei|last3=Zhang|first3=Lihua|last4=Yang|first4=Manyi|last5=Zhao|first5=Jinfeng|date=2019-02-25|title=Role of the novel gene BZW2 in the development of hepatocellular carcinoma|url=http://dx.doi.org/10.1002/jcp.28331|journal=Journal of Cellular Physiology|volume=234|issue=9|pages=16592–16600|doi=10.1002/jcp.28331|issn=0021-9541}}</ref><ref>{{Cite journal|last=Dong|first=Yuanmei|last2=Liu|first2=Yang|last3=Bai|first3=Hui|last4=Jiao|first4=Shunchang|date=2019-04-19|title=Systematic assessment of the clinicopathological prognostic significance of tissue cytokine expression for lung adenocarcinoma based on integrative analysis of TCGA data|url=http://dx.doi.org/10.1038/s41598-019-42345-0|journal=Scientific Reports|volume=9|issue=1|doi=10.1038/s41598-019-42345-0|issn=2045-2322}}</ref><ref>{{Cite journal|last=Gao|first=Haifeng|last2=Yu|first2=Guanghai|last3=Zhang|first3=Xian|last4=Yu|first4=Song|last5=Sun|first5=Yu|last6=Li|first6=Yinghua|date=2019-04|title=BZW2 gene knockdown induces cell growth inhibition, G1 arrest and apoptosis in muscle‐invasive bladder cancers: A microarray pathway analysis|url=http://dx.doi.org/10.1111/jcmm.14266|journal=Journal of Cellular and Molecular Medicine|volume=23|issue=6|pages=3905–3915|doi=10.1111/jcmm.14266|issn=1582-1838}}</ref> The AKT/mTOR pathway is an important intracellular signaling pathway that regulates the [[cell cycle]]. When the pathway activity is increased, cells proliferated at a higher rate and [[apoptosis]] decreases, leading to tumor growth. |
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=== SARS-CoV-2 === |
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BZW2 interacts with the nsp8 protein of [[Severe acute respiratory syndrome coronavirus 2|SARS-CoV-2]]. nsp8 dimerizes and forms a supercomplex which works to repress the translation of BZW2. |
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==References== |
==References== |
Revision as of 18:53, 31 July 2020
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Aliases | BZW2, MST017, MSTP017, HSPC028, basic leucine zipper and W2 domains 2, 5MP1 | ||||||||||||||||||||||||||||||||||||||||||||||||||
External IDs | MGI: 1914162 HomoloGene: 8530 GeneCards: BZW2 | ||||||||||||||||||||||||||||||||||||||||||||||||||
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Basic leucine zipper and W2 domain-containing protein 2 is a protein that is encoded by the BZW2 gene.[5][6] It is a eukaryotic translation factor and is conserved up to fungus, and further conserved in a couple bacteria species.
Gene
BZW2 is known as Basic Leucine Zipper W2 Domain-Containing Protein 2, MST017, MSTP017, 5MP1, and HSPC028.[7] It is located on chromosome 7 at the 21.1 locus. The gene spans 60,389 base pairs, at coordinates 16,583,248 – 16,804,999. There are 12 exons.
Protein
There are two known isoforms of BZW2. Isoform 1 is 419 amino acids long and is the most abundant form. Isoform 2 is 225 amino acids, containing only 11 exons and a shorter N-terminus.[7]
The coded protein is 419 amino acids long and weighs 48.3 kDa.[8] As described in the name, the protein contains a leucine-zipper motif. Four “L……” repeats are present in the beginning, giving rise to the characteristic leucine zipper helix within the 3D structure. An eIF5C domain follows the leucine motif.
BZW2 has a higher amount of Lysines and a lower amount of Prolines. There are no significant charge segments.[9]
The secondary structure contains a majority of alpha helices. The tertiary structure forms a repeated fold of alpha-helices, a structure that is conserved through bacteria.[10]
Regulation
Gene-Level
There are three known promoters for BZW2.[11] It is regulated by numerous transcription factors, including an estrogen receptor transcription factor, leucine zipper transcription factor, and Y sex-determining transcription factors. Overall BZW2 is ubiquitously expressed throughout tissues. There is increased mRNA abundance in the heart, placenta, and skeletal muscle.
Transcript-Level
There are four major stem loops in the 5’ untranslated and four in the 3’ untranslated region.[12]
Protein-Level
BZW2 is localized in the cytoplasm and has multiple phosphorylation, acetylation, glycosylation, SUMOylation and glycation sites for regulation.[13]
Evolution
BZW2 has a single paralog, BZW1 which is conserved up to plants.[14] There are BZW2 orthologs up to a couple species of bacteria. The most distant ortholog was Microbacterium aborescens. BZW2 contains an eIF5C domain which is also present in eIF2BE, eIF4G, eIF5, and a GAP protein specific for eIF2.[15]
Compared to Cytochrome C, a quickly diverging protein, and Fibronectin, a slowly diverging protein, BZW2 has had slow corrected divergence over time.
Interactions
BZW2 is known to interact with:
Clinical Significance
Cancer
BZW2 has been studied to determine its role in multiple cancers. Overall, the studies all showed that upregulation of BZW2 lead to more severe forms of cancer, higher rate of mortality, and increased likeliness of reoccurrence.
A 2019 study focused on the effect of BZW2 in colorectal cancer.[16] It found that upregulation of BZW2 promoted tumor growth and had a downstream upregulation effect on c-Myc, a proto-oncogene. A second study from 2020 determined this upregulation also had a positive effect on the activation of the ERK/MAPK pathway. [17]
In hepatocellular carcinoma, ostersarcoma, lung adenocarcinoma, and muscle-invasive bladder cancer, overexpression of BZW2 lead to overactivation of the AKT/mTOR signaling pathway by increasing phosphorylation of AKT and mTOR.[18][19][20][21] The AKT/mTOR pathway is an important intracellular signaling pathway that regulates the cell cycle. When the pathway activity is increased, cells proliferated at a higher rate and apoptosis decreases, leading to tumor growth.
SARS-CoV-2
BZW2 interacts with the nsp8 protein of SARS-CoV-2. nsp8 dimerizes and forms a supercomplex which works to repress the translation of BZW2.
References
- ^ a b c GRCh38: Ensembl release 89: ENSG00000136261 – Ensembl, May 2017
- ^ a b c GRCm38: Ensembl release 89: ENSMUSG00000020547 – Ensembl, May 2017
- ^ "Human PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.
- ^ "Mouse PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.
- ^ Zhang QH, Ye M, Wu XY, Ren SX, Zhao M, Zhao CJ, Fu G, Shen Y, Fan HY, Lu G, Zhong M, Xu XR, Han ZG, Zhang JW, Tao J, Huang QH, Zhou J, Hu GX, Gu J, Chen SJ, Chen Z (Nov 2000). "Cloning and functional analysis of cDNAs with open reading frames for 300 previously undefined genes expressed in CD34+ hematopoietic stem/progenitor cells". Genome Res. 10 (10): 1546–60. doi:10.1101/gr.140200. PMC 310934. PMID 11042152.
- ^ "Entrez Gene: BZW2 basic leucine zipper and W2 domains 2".
- ^ a b "BZW2 basic leucine zipper and W2 domains 2 [Homo sapiens (human)] - Gene - NCBI". www.ncbi.nlm.nih.gov. Retrieved 2020-07-31.
- ^ "BZW2 Antibody (PA5-21808)". www.thermofisher.com. Retrieved 2020-07-31.
- ^ Madeira, Fábio; Park, Young mi; Lee, Joon; Buso, Nicola; Gur, Tamer; Madhusoodanan, Nandana; Basutkar, Prasad; Tivey, Adrian R N; Potter, Simon C; Finn, Robert D; Lopez, Rodrigo (2019-04-12). "The EMBL-EBI search and sequence analysis tools APIs in 2019". Nucleic Acids Research. 47 (W1): W636–W641. doi:10.1093/nar/gkz268. ISSN 0305-1048.
- ^ Kelley, Lawrence A; Mezulis, Stefans; Yates, Christopher M; Wass, Mark N; Sternberg, Michael J E (2015-05-07). "The Phyre2 web portal for protein modeling, prediction and analysis". Nature Protocols. 10 (6): 845–858. doi:10.1038/nprot.2015.053. ISSN 1754-2189.
- ^ "Genomatix - NGS Data Analysis & Personalized Medicine". www.genomatix.de. Retrieved 2020-07-31.
- ^ Zuker, M. (2003-07-01). "Mfold web server for nucleic acid folding and hybridization prediction". Nucleic Acids Research. 31 (13): 3406–3415. doi:10.1093/nar/gkg595. ISSN 1362-4962.
- ^ "BZW2 (human)". www.phosphosite.org. Retrieved 2020-07-31.
- ^ "Protein BLAST: search protein databases using a protein query". blast.ncbi.nlm.nih.gov. Retrieved 2020-07-31.
- ^ Koonin, Eugene V. (1995-08). "Multidomain organization of eukaryotic guanine nucleotide exchange translation initiation factor eIF-2B subunits revealed by analysis of conserved sequence motifs". Protein Science. 4 (8): 1608–1617. doi:10.1002/pro.5560040819. ISSN 0961-8368.
{{cite journal}}
: Check date values in:|date=
(help) - ^ Sato, Kuniaki; Masuda, Takaaki; Hu, Qingjiang; Tobo, Taro; Gillaspie, Sarah; Niida, Atsushi; Thornton, Mackenzie; Kuroda, Yousuke; Eguchi, Hidetoshi; Nakagawa, Takashi; Asano, Katsura (2019-06). "Novel oncogene 5MP1 reprograms c-Myc translation initiation to drive malignant phenotypes in colorectal cancer". EBioMedicine. 44: 387–402. doi:10.1016/j.ebiom.2019.05.058. ISSN 2352-3964.
{{cite journal}}
: Check date values in:|date=
(help) - ^ Huang, Longping; Chen, Si; Fan, Haijun; Ai, Fan; Sheng, Weiwei (2019-10-23). "BZW2 promotes the malignant progression of colorectal cancer via activating the ERK/MAPK pathway". Journal of Cellular Physiology. 235 (5): 4834–4842. doi:10.1002/jcp.29361. ISSN 0021-9541.
- ^ Cheng, Dong-Dong; Li, Shi-Jie; Zhu, Bin; Yuan, Ting; Yang, Qing-Cheng; Fan, Cun-Yi (2017-04). "Downregulation of BZW2 inhibits osteosarcoma cell growth by inactivating the Akt/mTOR signaling pathway". Oncology Reports. 38 (4): 2116–2122. doi:10.3892/or.2017.5890. ISSN 1021-335X.
{{cite journal}}
: Check date values in:|date=
(help) - ^ Jin, Xin; Liao, Mingmei; Zhang, Lihua; Yang, Manyi; Zhao, Jinfeng (2019-02-25). "Role of the novel gene BZW2 in the development of hepatocellular carcinoma". Journal of Cellular Physiology. 234 (9): 16592–16600. doi:10.1002/jcp.28331. ISSN 0021-9541.
- ^ Dong, Yuanmei; Liu, Yang; Bai, Hui; Jiao, Shunchang (2019-04-19). "Systematic assessment of the clinicopathological prognostic significance of tissue cytokine expression for lung adenocarcinoma based on integrative analysis of TCGA data". Scientific Reports. 9 (1). doi:10.1038/s41598-019-42345-0. ISSN 2045-2322.
- ^ Gao, Haifeng; Yu, Guanghai; Zhang, Xian; Yu, Song; Sun, Yu; Li, Yinghua (2019-04). "BZW2 gene knockdown induces cell growth inhibition, G1 arrest and apoptosis in muscle‐invasive bladder cancers: A microarray pathway analysis". Journal of Cellular and Molecular Medicine. 23 (6): 3905–3915. doi:10.1111/jcmm.14266. ISSN 1582-1838.
{{cite journal}}
: Check date values in:|date=
(help)
External links
- Human BZW2 genome location and BZW2 gene details page in the UCSC Genome Browser.
Further reading
- Andersen JS, Lam YW, Leung AK, et al. (2005). "Nucleolar proteome dynamics". Nature. 433 (7021): 77–83. doi:10.1038/nature03207. PMID 15635413.
- Olsen JV, Blagoev B, Gnad F, et al. (2006). "Global, in vivo, and site-specific phosphorylation dynamics in signaling networks". Cell. 127 (3): 635–48. doi:10.1016/j.cell.2006.09.026. PMID 17081983.
- Ewing RM, Chu P, Elisma F, et al. (2007). "Large-scale mapping of human protein-protein interactions by mass spectrometry". Mol. Syst. Biol. 3 (1): 89. doi:10.1038/msb4100134. PMC 1847948. PMID 17353931.