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the 13th letter of the Roman alphabet (同)m

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Mach number / mark[s] / Monsieur

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出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2016/06/18 12:33:21」(JST)

Myc (c-Myc) is a regulator gene that codes for a transcription factor. The protein encoded by this gene is a multifunctional, nuclear phosphoprotein that plays a role in cell cycle progression, apoptosis and cellular transformation.^[1]

A mutated version of Myc is found in many cancers, which causes Myc to be constitutively (persistently) expressed. This leads to the unregulated expression of many genes, some of which are involved in cell proliferation, and results in the formation of cancer.^[1] A common human translocation involving Myc is critical to the development of most cases of Burkitt lymphoma.^[2] Malfunctions in Myc have also been found in carcinoma of the cervix, colon, breast, lung and stomach.^[1] Myc is thus viewed as a promising target for anti-cancer drugs.^[3]

In the human genome, Myc is located on chromosome 8 and is believed to regulate expression of 15% of all genes^[4] through binding on enhancer box sequences (E-boxes) and recruiting histone acetyltransferases (HATs). This means that in addition to its role as a classical transcription factor, Myc also functions to regulate global chromatin structure by regulating histone acetylation both in gene-rich regions and at sites far from any known gene.^[5]

Discovery

Myc gene was first discovered in Burkitt lymphoma patients. In Burkitt lymphoma, cancer cells show chromosomal translocations, in which chromosome 8 is frequently involved. Cloning the break-point of the fusion chromosomes revealed a gene that was similar to myelocytomatosis viral oncogene (v-Myc). Thus, the newfound cellular gene was named c-Myc.

Structure

Myc protein belongs to Myc family of transcription factors, which also includes N-Myc and L-Myc genes. Myc family of transcription factors contain bHLH/LZ (basic helix-loop-helix leucine zipper) domain. Myc protein, through its bHLH structural motif can bind to DNA, while the leucine zipper domain allows the dimerization with its partner Max, another bHLH transcription factor.

Myc mRNA contains an IRES (internal ribosome entry site) that allows the RNA to be translated into protein when 5' cap-dependent translation is inhibited, such as during viral infection.

Function

Myc protein is a transcription factor that activates expression of many genes through binding enhancer box sequences (E-boxes) and recruiting histone acetyltransferases (HATs). It can also act as a transcriptional repressor. By binding Miz-1 transcription factor and displacing the p300 co-activator, it inhibits expression of Miz-1 target genes. In addition, myc has a direct role in the control of DNA replication.^[6]

Myc is activated upon various mitogenic signals such as Wnt, Shh and EGF (via the MAPK/ERK pathway). By modifying the expression of its target genes, Myc activation results in numerous biological effects. The first to be discovered was its capability to drive cell proliferation (upregulates cyclins, downregulates p21), but it also plays a very important role in regulating cell growth (upregulates ribosomal RNA and proteins), apoptosis (downregulates Bcl-2), differentiation, and stem cell self-renewal. Myc is a very strong proto-oncogene and it is very often found to be upregulated in many types of cancers. Myc overexpression stimulates gene amplification,^[7] presumably through DNA over-replication.

There have been several studies that have clearly indicated Myc's role in cell competition.^[8]

A major effect of Myc is B cell proliferation.^[9]

c-Myc induces MTDH(AEG-1) gene expression and in turn itself requires AEG-1 oncogene for its expression.

Myc-nick

Myc-nick is a cytoplasmic form of Myc produced by a partial proteolytic cleavage of full-length c-Myc and N-Myc.^[10] Myc cleavage is mediated by the calpain family of calcium-dependent cytosolic proteases.

The cleavage of Myc by calpains is a constitutive process but is enhanced under conditions that require rapid downregulation of Myc levels, such as during terminal differentiation. Upon cleavage, the C-terminus of Myc (containing the DNA binding domain) is degraded, while Myc-nick, the N-terminal segment 298-residue segment remains in the cytoplasm. Myc-nick contains binding domains for histone acetyltransferases and for ubiquitin ligases.

The functions of Myc-nick are currently under investigation, but this new Myc family member was found to regulate cell morphology, at least in part, by interacting with acetyl transferases to promote the acetylation of α-tubulin. Ectopic expression of Myc-nick accelerates the differentiation of committed myoblasts into muscle cells.

Clinical significance

Except for early response genes, Myc universally upregulates gene expression. Furthermore, the upregulation is nonlinear. Genes whose expression is already significantly upregulated in the absence of Myc are strongly boosted in the presence of Myc, whereas genes whose expression is low in the absence Myc get only a small boost when Myc is present.^[11]

Inactivation of SUMO-activating enzyme (SAE1 / SAE2) in the presence of Myc hyperactivation results in mitotic catastrophe and cell death in cancer cells. Hence inhibitors of SUMOylation may be a possible treatment for cancer.^[12]

Amplification of the MYC gene was found in a significant number of epithelial ovarian cancer cases.^[13] In TCGA datasets, the amplification of Myc occurs in several cancer types, including breast, colorectal, pancreatic, gastric, and uterine cancers.^[14]

In the experimental transformation process of normal cells into cancer cells, the MYC gene can cooperate with the RAS gene.^[15]^[16]

Expression of Myc is highly dependent on BRD4 function in some cancers.^[17]^[18] BET inhibitors have been used to successfully block Myc function in pre-clinical cancer models and are currently being evaluated in clinical trials.^[19]^[20]

Animal Models

During the discovery of Myc gene, it was realized that chromosomes that reciprocally translocate to chromosome 8 contained immunoglobulin genes at the break-point. Enhancers that normally drive expression of immunoglobin genes now lead to overexpression of Myc proto-oncogene in lymphoma cells. To study the mechanism of tumorigenesis in Burkitt lymphoma by mimicking expression pattern of Myc in these cancer cells, transgenic mouse models were developed. Myc gene placed under the control of IgM heavy chain enhancer in transgenic mice gives rise to mainly lymphomas. Later on, in order to study effects of Myc in other types of cancer, transgenic mice that overexpress Myc in different tissues (liver, breast) were also made. In all these mouse models overexpression of Myc causes tumorigenesis, illustrating the potency of Myc oncogene. In a study with mice, reduced expression of Myc was shown to induce longevity, with significantly extended median and maximum lifespans in both sexes and a reduced mortality rate across all ages, better health, cancer progression was slower, better metabolism and they had smaller bodies. also, Less TOR, AKT, S6K and other changes in energy and metabolic pathways (such as AMPK, more oxygen consumption, more body movements, etc). The study by John M. Sedivy and others used Cre-Loxp -recombinase to knockout one copy of Myc and this resulted in a "Haplo-insufficient" genotype noted as Myc+/-. The phenotypes seen oppose the effects of normal aging and are shared with many other long-lived mouse models such as CR (calorie restriction) ames dwarf, rapamycin, metformin and resveratrol. One study found that Myc and p53 genes were key to the survival of Chronic Myeloid Leukaemia (CML) cells. Targeting Myc and p53 proteins with drugs gave positive results on mice with CML.^[21]^[22]

Interactions

Myc has been shown to interact with:

ACTL6A^[23]
BRCA1^[24]^[25]^[26]^[27]
Bcl-2^[28]
Cyclin T1^[29]
CHD8.^[30]
DNMT3A^[31]
EP400^[32]
GTF2I^[33]
HTATIP^[34]
let-7^[35]^[36]^[37]
MAPK1^[28]^[38]^[39]
MAPK8^[40]
MAX^[41]^[42]^[43]^[44]^[45]^[46]^[47]^[48]^[49]^[50]^[51]^[52]^[53]
MLH1^[45]
MYCBP2^[54]
MYCBP^[55]
NMI^[24]
NFYB^[56]
NFYC^[57]
P73^[58]
PCAF^[59]
PFDN5^[60]^[61]
RuvB-like 1^[23]^[32]
SAP130^[59]
SMAD2^[62]
SMAD3^[62]
SMARCA4^[23]^[41]
SMARCB1^[44]
SUPT3H^[59]
TIAM1^[63]
TADA2L^[59]
TAF9^[59]
TFAP2A^[64]
TRRAP^[23]^[42]^[43]^[59]
WDR5^[65]
YY1^[66] and
ZBTB17.^[67]^[68]

Overview of signal transduction pathways involved in apoptosis.

References

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External links

The Myc Protein
NCBI Human Myc protein
Myc cancer gene
myc Proto-Oncogene Proteins at the US National Library of Medicine Medical Subject Headings (MeSH)
Generating iPS Cells from MEFS through Forced Expression of Sox-2, Oct-4, c-Myc, and Klf4
Drosophila Myc - The Interactive Fly
FactorBook C-Myc

UpToDate Contents

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1. バーキットリンパ腫の病理学 pathobiology of burkitt lymphoma
2. びまん性大細胞型B細胞性リンパ腫および原発性縦隔大細胞型B細胞リンパ腫の病理学 pathobiology of diffuse large b cell lymphoma and primary mediastinal large b cell lymphoma
3. 神経芽腫の疫学、病因、および病理 epidemiology pathogenesis and pathology of neuroblastoma
4. 甲状腺結節および非髄様甲状腺癌におけるオンコジーンおよび腫瘍抑制遺伝子 oncogenes and tumor suppressor genes in thyroid nodules and nonmedullary thyroid cancer
5. 大腸癌の分子遺伝学 molecular genetics of colorectal cancer

English Journal

Combinational delivery of c-myc siRNA and nucleoside analogs in a single, synthetic nanocarrier for targeted cancer therapy.

Zhang Y, Peng L, Mumper RJ, Huang L.SourceDivision of Molecular Pharmaceutics, Center for Nanotechnology in Drug Delivery, Eshelman School of Pharmacy, University of North Carolina at Chapel Hill, Chapel Hill, NC 27599, USA.
Biomaterials.Biomaterials.2013 Nov;34(33):8459-68. doi: 10.1016/j.biomaterials.2013.07.050. Epub 2013 Aug 8.
The treatment of aggressive non-small-cell lung cancer (NSCLC) depends on the creation of new therapeutic regimens in clinical settings. In this study, we developed a Lipid/Calcium/Phosphate (LCP) nanoparticle that combines chemotherapy with gene therapy. By encapsulating a chemodrug, gemcitabine mo
PMID 23932296

N-Myc knockdown and apigenin treatment controlled growth of malignant neuroblastoma cells having N-Myc amplification.

Hossain MM, Banik NL, Ray SK.SourceUniversity of South Carolina School of Medicine, Department of Pathology, Microbiology, and Immunology, Columbia, SC 29209, USA.
Gene.Gene.2013 Oct 15;529(1):27-36. doi: 10.1016/j.gene.2013.07.094. Epub 2013 Aug 11.
Malignant neuroblastomas mostly occur in children and are frequently associated with N-Myc amplification. Oncogene amplification, which is selective increase in copy number of the oncogene, provides survival advantages in solid tumors including malignant neuroblastoma. We have decreased expression o
PMID 23941992

Japanese Journal

Recurrent amplification of MYC and TNFRSF11B in 8q24 is associated with poor survival in patients with gastric cancer

Gastric cancer : official journal of the International Gastric Cancer Association and the Japanese Gastric Cancer Association 19(1), 116-127, 2016
NAID 40020700056

Evaluation of phenotypic, functional and molecular characteristics of porcine mesenchymal stromal/stem cells depending on donor age, gender and tissue source

Journal of Veterinary Medical Science advpub(0), 2016
NAID 130005127215

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★リンクテーブル★

国試過去問	「105G032」
リンク元	「14番染色体」「8番染色体」「t(8;14)」
関連記事	「M」

「105G032」

MYC
Available structures
PDB	Ortholog search: PDBe RCSB
List of PDB id codes
	1A93, 1EE4, 1MV0, 1NKP, 2A93, 2OR9, 4Y7R
Identifiers
Aliases	MYC, MRTL, MYCC, bHLHe39, c-Myc, v-myc avian myelocytomatosis viral oncogene homolog
External IDs	MGI: 97250 HomoloGene: 31092 GeneCards: 4609
Gene ontology
Molecular function	• DNA binding; • protein dimerization activity; • transcription factor activity, sequence-specific DNA binding; • protein complex binding; • transcriptional activator activity, RNA polymerase II core promoter proximal region sequence-specific binding; • transcription factor binding; • RNA polymerase II core promoter proximal region sequence-specific DNA binding; • E-box binding; • protein binding; • repressing transcription factor binding;
Cellular component	• cytosol; • nucleoplasm; • protein complex; • nucleolus; • nucleus;
Biological process	• Notch signaling pathway; • chromatin remodeling; • negative regulation of monocyte differentiation; • positive regulation of metanephric cap mesenchymal cell proliferation; • regulation of transcription, DNA-templated; • gene expression; • regulation of telomere maintenance; • positive regulation of epithelial cell proliferation; • positive regulation of fibroblast proliferation; • cellular response to UV; • oxygen transport; • negative regulation of apoptotic process; • negative regulation of transcription from RNA polymerase II promoter; • chromosome organization; • transcription, DNA-templated; • MAPK cascade; • cellular response to DNA damage stimulus; • positive regulation of transcription, DNA-templated; • positive regulation of cysteine-type endopeptidase activity involved in apoptotic process; • branching involved in ureteric bud morphogenesis; • cellular response to drug; • negative regulation of cell division; • fibroblast apoptotic process; • positive regulation of mesenchymal cell proliferation; • cell cycle arrest; • positive regulation of response to DNA damage stimulus; • positive regulation of cell proliferation; • positive regulation of DNA biosynthetic process; • regulation of gene expression; • canonical Wnt signaling pathway; • negative regulation of stress-activated MAPK cascade; • energy reserve metabolic process; • response to growth factor; • transcription initiation from RNA polymerase II promoter; • transforming growth factor beta receptor signaling pathway; • response to gamma radiation; • negative regulation of fibroblast proliferation; • response to drug; • positive regulation of transcription from RNA polymerase II promoter; • cellular iron ion homeostasis;
	Sources:Amigo / QuickGO
Orthologs
	4609
	17869
	ENSG00000136997
	ENSMUSG00000022346
	P01106
	P01108
	NM_002467
	NM_001177352; NM_001177353; NM_001177354; NM_010849
	NP_002458.2
	NP_001170823.1; NP_001170824.1; NP_001170825.1
	Wikidata
View/Edit Human	View/Edit Mouse

　　[★]

機能の不活性化がヒトでの発癌につながる遺伝子はどれか。 2つ選べ。

a　c-myc(MYC)
b　erbB2(ERBB2)
c　K-ras(KRAS)
d　p53(TP53)
e　Rb(RB)

[正答]

※国試ナビ4※　［105G031］←［国試_105］→［105G033］

「14番染色体」

　　[★]

英: chromosome 14
同: 第14番染色体、第14染色体

t(8;14)(q24;q32)［MYC /IgH］バーキットリンパ腫
t(14;18)［IGH /BCL2］濾胞リンパ腫

「8番染色体」

　　[★]

英: chromosome 8
同: 第8番染色体、第8染色体

t(8;14)(q24;q32)［MYC /IgH］バーキットリンパ腫
t(8;21)[AML1;ETO]　AML-M2

「t(8;14)」

　　[★]

t(8;14)(q24;q32)［MYC;IgH］バーキットリンパ腫

「M」

　　[★] メチオニン methionine 　　

[entrez-1] "Myc". NCBI.

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