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Proto-Oncogenes

Genetics and Genomics · NLM MeSH · updated 2026-10-29 · version 1

Citations verified against PubMedNot yet clinician-reviewed10 referencesApproximately 4 min read

Normal cellular genes homologous to viral oncogenes.

Table of Contents

1. Definition2. Research landscape3. Reading the evidence4. Open-access studies5. Key reviews6. References7. About this article

Definition and research guide

This page combines a MeSH definition, publication statistics and selected research abstracts. It is a starting point for reading the literature, not a medical textbook or a systematic evidence review.

Content available: 4 complete abstracts (835 words), 10 references. Full article text is not stored or reproduced here; follow the PMID, DOI or PMC links to read the original sources and their access terms.

Definition

Normal cellular genes homologous to viral oncogenes. The products of proto-oncogenes are important regulators of biological processes and appear to be involved in the events that serve to maintain the ordered procession through the cell cycle. Proto-oncogenes have names of the form c-onc.

Definition: NLM Medical Subject Headings (MeSH) scope note (Public domain (U.S. National Library of Medicine); attribution requested). View the MeSH record.

Also known as: Proto Oncogenes; Proto-Oncogene; Proto Oncogene.

Where it fits in the MeSH classification

Broader topics
Oncogenes
Narrower topics
Genes, abl; Genes, bcl-1; Genes, bcl-2; Genes, erbA; Genes, erbB; Genes, fms; Genes, fos; Genes, jun; Genes, mos; Genes, myb; Genes, myc; Genes, ras; Genes, rel; Genes, sis

Research landscape

PubMed indexes 41,882 records tagged with the MeSH heading “Proto-Oncogenes” (counts retrieved 2026-10-29). 1,437 of them (3.4%) were published from 2020 onward, and the busiest period so far is through 1999 with 20,162 records.

Reviews account for 4,981 records (11.9%) and meta-analyses for 82. Randomized controlled trials number 168 (0.4%) as a share of all records on the topic. Randomized trials make up only a small share of this literature, so much of what is known comes from laboratory, observational or descriptive work.

PubMed records by publication periodthrough 1999: 20,162 records; 2000-2009: 13,329 records; 2010-2019: 6,954 records; 2020 onward: 1,437 records20,162through 199913,3292000-20096,9542010-20191,4372020 onward
Records in PubMed indexed with this heading, by publication period. Real counts from NCBI E-utilities; the final period covers only part of the current year. Volume reflects how much has been published, not how strong the evidence is.
Show the data
PeriodRecords
through 199920,162
2000-200913,329
2010-20196,954
2020 onward1,437
Share of records by publication typeReviews: 11.9 percent; Meta-analyses: 0.2 percent; Randomized controlled trials: 0.4 percentReviews11.9% (4,981)Meta-analyses0.2% (82)Randomized controlled trials0.4% (168)
Share of all PubMed records on this heading that carry each publication type. Types overlap with other records, so the shares do not sum to 100%.
Publication-type chart data
Publication types among 41,882 PubMed records
TypeRecordsShare
Reviews4,98111.9%
Meta-analyses820.2%
Randomized controlled trials1680.4%

Method: PubMed query "Proto-Oncogenes"[MeSH Terms] with publication-type and date filters, retrieved 2026-10-29.

Reading the evidence in genetics and genomics

Genetic findings vary widely in strength. A variant linked to a condition in one study may not replicate in other ancestries, and most common variants have small effects that are meaningful only in aggregate. Studies of rare, high-impact variants can support strong causal claims about a gene, while association studies describe statistical links that need functional and independent replication. Check the sample size, the ancestry of participants, whether results were replicated, and whether a laboratory or clinical outcome was actually measured.

Evidence from open-access studies

These open-access studies come from PubMed’s best-match results for this heading (papers whose main topic it is), ranked here by citation count, with one recent addition. Each abstract is reproduced in full under its recorded open licence so that you can read the authors’ own words. They illustrate the literature; they are not a systematic summary of it. Citation counts measure attention, not reliability or clinical benefit.

Study comparison — selected abstracts, not a systematic review
Study / authorsYearSource label / designEurope PMC citationsLicence / source
Deletion of a Csf1r enhancer selectively impacts CSF1R expression and development of tissue macrophage populations
Rojo R, Raper A, Ozdemir DD, Lefevre L, Grabert K, Wollscheid-Lengeling E, et al.
2019Research article
Design not recorded in structured metadata
350cc by
PMID 31324781 · DOI · PMC6642117
c-MYC mediates the crosstalk between breast cancer cells and tumor microenvironment
Gao FY, Li XT, Xu K, Wang RT, Guan XX
2023Review article
Design not recorded in structured metadata
152cc by
PMID 36721232 · DOI · PMC9887805
Acute depletion of CTCF directly affects MYC regulation through loss of enhancer-promoter looping
Hyle J, Zhang Y, Wright S, Xu B, Shao Y, Easton J, et al.
2019Research article
Design not recorded in structured metadata
136cc by
PMID 31127282 · DOI · PMC6648894
Mitotic gene regulation by the N-MYC-WDR5-PDPK1 nexus
Streeter SA, Williams AG, Evans JR, Wang J, Guarnaccia AD, Florian AC, et al.
2024Research article
Design not recorded in structured metadata
1cc by
PMID 38605297 · DOI · PMC11007937

Deletion of a Csf1r enhancer selectively impacts CSF1R expression and development of tissue macrophage populations [1]

Research article Rojo R, Raper A, Ozdemir DD, Lefevre L, Grabert K, Wollscheid-Lengeling E, et al. · · 2019 · cited 350 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
The proliferation, differentiation and survival of mononuclear phagocytes depend on signals from the receptor for macrophage colony-stimulating factor, CSF1R.
Closing excerpt
Csf1rΔFIRE/ΔFIRE mice thus provide a model to explore the homeostatic, physiological and immunological functions of tissue-specific macrophage populations in adult animals.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (144 words)

The proliferation, differentiation and survival of mononuclear phagocytes depend on signals from the receptor for macrophage colony-stimulating factor, CSF1R. The mammalian Csf1r locus contains a highly conserved super-enhancer, the fms-intronic regulatory element (FIRE). Here we show that genomic deletion of FIRE in mice selectively impacts CSF1R expression and tissue macrophage development in specific tissues. Deletion of FIRE ablates macrophage development from murine embryonic stem cells. Csf1rΔFIRE/ΔFIRE mice lack macrophages in the embryo, brain microglia and resident macrophages in the skin, kidney, heart and peritoneum. The homeostasis of other macrophage populations and monocytes is unaffected, but monocytes and their progenitors in bone marrow lack surface CSF1R. Finally, Csf1rΔFIRE/ΔFIRE mice are healthy and fertile without the growth, neurological or developmental abnormalities reported in Csf1r-/- rodents. Csf1rΔFIRE/ΔFIRE mice thus provide a model to explore the homeostatic, physiological and immunological functions of tissue-specific macrophage populations in adult animals.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 31324781 · DOI · PMC6642117

c-MYC mediates the crosstalk between breast cancer cells and tumor microenvironment [2]

Review article Gao FY, Li XT, Xu K, Wang RT, Guan XX · · 2023 · cited 152 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
The MYC oncogenic family is dysregulated in diverse tumors which is generally linked to the poor prognosis of tumors.
Closing excerpt
Video Abstract.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (169 words)

The MYC oncogenic family is dysregulated in diverse tumors which is generally linked to the poor prognosis of tumors. The members in MYC family are transcription factors which are responsible for the regulation of various genes expression. Among them, c-MYC is closely related to the progression of tumors. Furthermore, c-MYC aberrations is tightly associated with the prevalence of breast cancer. Tumor microenvironment (TME) is composed of many different types of cellular and non-cellular factors, mainly including cancer-associated fibroblasts, tumor-associated macrophages, vascular endothelial cells, myeloid-derived suppressor cells and immune cells, all of which can affect the diagnosis, prognosis, and therapeutic efficacy of breast cancer. Importantly, the biological processes occurred in TME, such as angiogenesis, immune evasion, invasion, migration, and the recruition of stromal and tumor-infiltrating cells are under the modulation of c-MYC. These findings indicated that c-MYC serves as a critical regulator of TME. Here, we aimed to summarize and review the relevant research, thus to clarify c-MYC is a key mediator between breast cancer cells and TME. Video Abstract.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 36721232 · DOI · PMC9887805

Acute depletion of CTCF directly affects MYC regulation through loss of enhancer-promoter looping [3]

Research article Hyle J, Zhang Y, Wright S, Xu B, Shao Y, Easton J, et al. · · 2019 · cited 136 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
Numerous pieces of evidence support the complex, 3D spatial organization of the genome dictates gene expression.
Closing excerpt
Notably, MYC expression was not profoundly affected upon CTCF loss in HUDEP-2 cells suggesting that CTCF could play a B-ALL cell line specific role in maintaining MYC expression.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (197 words)

Numerous pieces of evidence support the complex, 3D spatial organization of the genome dictates gene expression. CTCF is essential to define topologically associated domain boundaries and to facilitate the formation of insulated chromatin loop structures. To understand CTCF's direct role in global transcriptional regulation, we integrated the miniAID-mClover3 cassette to the endogenous CTCF locus in a human pediatric B-ALL cell line, SEM, and an immortal erythroid precursor cell line, HUDEP-2, to allow for acute depletion of CTCF protein by the auxin-inducible degron system. In SEM cells, CTCF loss notably disrupted intra-TAD loops and TAD integrity in concurrence with a reduction in CTCF-binding affinity, while showing no perturbation to nuclear compartment integrity. Strikingly, the overall effect of CTCF's loss on transcription was minimal. Whole transcriptome analysis showed hundreds of genes differentially expressed in CTCF-depleted cells, among which MYC and a number of MYC target genes were specifically downregulated. Mechanically, acute depletion of CTCF disrupted the direct interaction between the MYC promoter and its distal enhancer cluster residing ∼1.8 Mb downstream. Notably, MYC expression was not profoundly affected upon CTCF loss in HUDEP-2 cells suggesting that CTCF could play a B-ALL cell line specific role in maintaining MYC expression.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 31127282 · DOI · PMC6648894

Mitotic gene regulation by the N-MYC-WDR5-PDPK1 nexus [4]

Research article Streeter SA, Williams AG, Evans JR, Wang J, Guarnaccia AD, Florian AC, et al. · · 2024 · cited 1 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
BackgroundDuring mitosis the cell depends on proper attachment and segregation of replicated chromosomes to generate two identical progeny.
Closing excerpt
This study provides a foundation for future studies to determine the exact mechanism by which N-MYC, WDR5, and PDPK1 converge on cell cycle related processes.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (325 words)

BackgroundDuring mitosis the cell depends on proper attachment and segregation of replicated chromosomes to generate two identical progeny. In cancers defined by overexpression or dysregulation of the MYC oncogene this process becomes impaired, leading to genomic instability and tumor evolution. Recently it was discovered that the chromatin regulator WDR5-a critical MYC cofactor-regulates expression of genes needed in mitosis through a direct interaction with the master kinase PDPK1. However, whether PDPK1 and WDR5 contribute to similar mitotic gene regulation in MYC-overexpressing cancers remains unclear. Therefore, to characterize the influence of WDR5 and PDPK1 on mitotic gene expression in cells with high MYC levels, we performed a comparative transcriptomic analysis in neuroblastoma cell lines defined by MYCN-amplification, which results in high cellular levels of the N-MYC protein.ResultsUsing RNA-seq analysis, we identify the genes regulated by N-MYC and PDPK1 in multiple engineered CHP-134 neuroblastoma cell lines and compare them to previously published gene expression data collected in CHP-134 cells following inhibition of WDR5. We find that as expected N-MYC regulates a multitude of genes, including those related to mitosis, but that PDPK1 regulates specific sets of genes involved in development, signaling, and mitosis. Analysis of N-MYC- and PDPK1-regulated genes reveals a small group of commonly controlled genes associated with spindle pole formation and chromosome segregation, which overlap with genes that are also regulated by WDR5. We also find that N-MYC physically interacts with PDPK1 through the WDR5-PDPK1 interaction suggesting regulation of mitotic gene expression may be achieved through a N-MYC-WDR5-PDPK1 nexus.ConclusionsOverall, we identify a small group of genes highly enriched within functional gene categories related to mitotic processes that are commonly regulated by N-MYC, WDR5, and PDPK1 and suggest that a tripartite interaction between the three regulators may be responsible for setting the level of mitotic gene regulation in N-MYC amplified cell lines. This study provides a foundation for future studies to determine the exact mechanism by which N-MYC, WDR5, and PDPK1 converge on cell cycle related processes.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 38605297 · DOI · PMC11007937

Key reviews and meta-analyses

The most cited review-type papers among PubMed’s best-match results for this heading, listed with bibliographic details only. Follow the links for the full text and access terms.

  • Research article RAS oncogenes: weaving a tumorigenic web [5]
    Pylayeva-Gupta Y, Grabocka E, Grabocka E, Bar-Sagi D · Nat Rev Cancer · 2011 · cited 1,461 times (Europe PMC)
  • Research article Reflecting on 25 years with MYC [6]
    Meyer N, Penn LZ · Nat Rev Cancer · 2008 · cited 1,295 times (Europe PMC)
  • Research article Hyperactive Ras in developmental disorders and cancer [7]
    Schubbert S, Shannon K, Bollag G · Nat Rev Cancer · 2007 · cited 1,173 times (Europe PMC)
  • Research article Ras oncogenes: split personalities [8]
    Karnoub AE, Weinberg RA · Nat Rev Mol Cell Biol · 2008 · cited 1,155 times (Europe PMC)
  • Research article The oncogene HER2: its signaling and transforming functions and its role in human cancer pathogenesis [9]
    Moasser MM · Oncogene · 2007 · cited 917 times (Europe PMC)
  • Research article The MYC oncogene - the grand orchestrator of cancer growth and immune evasion [10]
    Dhanasekaran R, Deutzmann A, Mahauad-Fernandez WD, Hansen AS, Gouw AM, Felsher DW · Nat Rev Clin Oncol · 2022 · cited 876 times (Europe PMC)

References

  1. Rojo R, Raper A, Ozdemir DD, Lefevre L, Grabert K, Wollscheid-Lengeling E, et al.. Deletion of a Csf1r enhancer selectively impacts CSF1R expression and development of tissue macrophage populations. . 2019. PMID 31324781 · DOI · PMC6642117
  2. Gao FY, Li XT, Xu K, Wang RT, Guan XX. c-MYC mediates the crosstalk between breast cancer cells and tumor microenvironment. . 2023. PMID 36721232 · DOI · PMC9887805
  3. Hyle J, Zhang Y, Wright S, Xu B, Shao Y, Easton J, et al.. Acute depletion of CTCF directly affects MYC regulation through loss of enhancer-promoter looping. . 2019. PMID 31127282 · DOI · PMC6648894
  4. Streeter SA, Williams AG, Evans JR, Wang J, Guarnaccia AD, Florian AC, et al.. Mitotic gene regulation by the N-MYC-WDR5-PDPK1 nexus. . 2024. PMID 38605297 · DOI · PMC11007937
  5. Pylayeva-Gupta Y, Grabocka E, Grabocka E, Bar-Sagi D. RAS oncogenes: weaving a tumorigenic web. Nat Rev Cancer. 2011. PMID 21993244 · DOI · PMC3632399
  6. Meyer N, Penn LZ. Reflecting on 25 years with MYC. Nat Rev Cancer. 2008. PMID 19029958 · DOI
  7. Schubbert S, Shannon K, Bollag G. Hyperactive Ras in developmental disorders and cancer. Nat Rev Cancer. 2007. PMID 17384584 · DOI
  8. Karnoub AE, Weinberg RA. Ras oncogenes: split personalities. Nat Rev Mol Cell Biol. 2008. PMID 18568040 · DOI · PMC3915522
  9. Moasser MM. The oncogene HER2: its signaling and transforming functions and its role in human cancer pathogenesis. Oncogene. 2007. PMID 17471238 · DOI · PMC3021475
  10. Dhanasekaran R, Deutzmann A, Mahauad-Fernandez WD, Hansen AS, Gouw AM, Felsher DW. The MYC oncogene - the grand orchestrator of cancer growth and immune evasion. Nat Rev Clin Oncol. 2022. PMID 34508258 · DOI · PMC9083341

About this article

Sources
NLM Medical Subject Headings (definition, synonyms); NCBI PubMed (counts, identifiers); Europe PMC (open-access abstracts and citation counts).
Verification
Every reference was resolved against PubMed on 2026-10-31; title, year and DOI matched.
Review state
automated source verified; no human expert review has been recorded yet.
Version
1, last updated 2026-10-29.

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At a glance

41,882
PubMed records on this heading
3%
published since 2020
4,981
review articles
168
randomized trials
10
verified references below

Counts come from PubMed on 2026-10-29; they show how much has been written, not how strong the evidence is.

Related topics See all →

  • OncogenesGenetics and Genomics
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  • CodonGenetics and Genomics
  • EpigenomeGenetics and Genomics

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