FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Human Disease Model Report: cancer, multiple, Notch signaling pathway
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General Information
Name
cancer, multiple, Notch signaling pathway
FlyBase ID
FBhh0000766
Disease Ontology Term
Parent Disease
OMIM
Overview

Notch signaling is an evolutionarily conserved intercellular signaling pathway that participates in critical processes throughout development, from embryogenesis to determination of adult stem cells. Activation of the Notch signaling pathway is also observed in a number of cancers. Many of the genes in this pathway were first identified in Drosophila; experiments in flies continue to contribute to understanding of the Notch signaling pathway, including its role in development and progression of cancer.

In human, there are 4 genes that encode NOTCH family transmembrane receptor proteins (NOTCH1-NOTCH4); there are 2 genes similar to the Drosophila Notch ligand Delta (DLL1 and DLL4) and 2 genes similar to the Drosophila Notch ligand Serrate (JAG1 and JAG2).

Much of the work in flies investigating the role of the Notch pathway in development of cancer has involved combining Notch signaling dysregulation with another cancer model (see 'Related Diseases' below). This is typically achieved by using targeted expression of an activated form of Notch (N) or overexpression of one of the Notch ligands, Delta (Dl) or Serrate (Ser). Frequently used are constructs of Notch lacking its extracellular domain; the functional intracellular domain (NICD) is constitutively active. Genes potentially involved in tumorigenesis have also been identified using these systems. For example, overexpression of Dl in the adult eye results in a mild overgrowth phenotype (often called a 'sensitized model"); mutations of interacting genes have been identified that enhance this phenotype and result in much increased, tumorous overgrowth.

The loss of N, Dl or other Notch pathway components in stem cells results in a population of differentiation-defective cells; this system has been used to investigate stem-cell-derived cancers, such as intestinal stem cell tumors and neural stem cell tumors. See human disease model reports 'cancer, intestinal stem cell models' (FBhh0000767) and 'cancer, neural stem cell dedifferentiation' (FBhh0001139).

[updated Feb. 2022 by FlyBase; FBrf0222196]

Disease Summary Information
Disease Summary: cancer, multiple, Notch signaling pathway
OMIM report
Human gene(s) implicated
Symptoms and phenotype

The Notch signaling pathway is commonly activated in cancer and its activation plays key roles in cancer development and progression. Inappropriate Notch signalling has been associated with virtually all aspects of cancer, including tumor cell growth, survival, apoptosis, angiogenesis, invasion and metastasis (Dominguez, 2014; pubmed:24780858, FBrf0225143).

(FlyBase Curators, 2013-)
Genetics
Cellular phenotype and pathology
Molecular information

Notch signaling is an intercellular signaling pathway that regulates interactions between physically adjacent cells through binding of Notch family receptors to their membrane-bound ligands on the adjacent cells. The NOTCH family of proteins are Type I single-pass transmembrane proteins that share structural characteristics including an extracellular domain consisting of multiple epidermal growth factor-like (EGF) repeats, and an intracellular domain consisting of multiple different domain types. [from Gene Cards, NOTCH1; 2018.03.15]

(FlyBase Curators, 2013-)
External links
Disease synonyms
Search term: hyperplastic phenotype(s)
Ortholog Information
Human gene(s) in FlyBase
Other mammalian ortholog(s) used
    D. melanogaster Gene Information (3)
    Dmel\N
    Gene Snapshot
    Contributions welcome
    Molecular function (GO)
    Cellular component (GO)
    Gene Groups / Pathways
    Comments on ortholog(s)

    Moderate- to high-scoring ortholog of human NOTCH1, NOTCH2, NOTCH3, NOTCH4 (1 Drosophila to 4 human). Dmel\N shares 43-44% identity and 56-57% similarity with NOTCH1 and NOTCH2.

    Orthologs and Alignments from DRSC
    DIOPT - DRSC Integrative Ortholog Prediction Tool - Click the link below to search for orthologs in Humans
    H. sapiens (Human)
    Dmel\Delta
    Gene Snapshot
    Delta (Delta) encodes a single pass transmembrane EGF family protein and one of two ligands of the Notch signaling pathway. It regulates cell fate decisions and cell proliferation. Post transcriptional modification (such as by fucosylation, ubiquitination and proteolysis) of the product of Delta is key for its functions. [Date last reviewed: 2019-03-07]
    Molecular function (GO)
    Cellular component (GO)
    Gene Groups / Pathways
    Comments on ortholog(s)

    Moderate- to high-scoring ortholog of human DLL1 and DLL4 (1 Drosophila to 2 human); additional less closely related genes in human. Dmel\Dl shares 39% identity and 51-53% similarity with human DLL1 and DLL4.

    Orthologs and Alignments from DRSC
    DIOPT - DRSC Integrative Ortholog Prediction Tool - Click the link below to search for orthologs in Humans
    H. sapiens (Human)
    Dmel\Ser
    Gene Snapshot
    Contributions welcome
    Molecular function (GO)
    Cellular component (GO)
    Gene Groups / Pathways
    Comments on ortholog(s)

    Moderate-scoring ortholog of human JAG1 and JAG2 (1 Drosophila to 2 human). Dmel\Ser shares 33-35% identity and 46% similarity with the human genes.

    Orthologs and Alignments from DRSC
    DIOPT - DRSC Integrative Ortholog Prediction Tool - Click the link below to search for orthologs in Humans
    H. sapiens (Human)
    Other Genes Used: Viral, Bacterial, Synthetic (0)
      Summary of Physical Interactions (69 groups)
      N
      protein-protein
      Interacting group
      Assay
      References
      N
      two hybrid, enzyme linked immunosorbent assay, electron microscopy, molecular weight
      (Kelly et al., 2010, Pei and Baker, 2008, Fortini and Artavanis-Tsakonas, 1994)
      N - Abl
      anti bait coimmunoprecipitation, western blot
      (Koca et al., 2022)
      N - Akap200
      anti tag coimmunoprecipitation, western blot, anti tag western blot
      (Bala Tannan et al., 2018)
      N - arm
      anti bait coimmunoprecipitation, western blot
      (Koca et al., 2022, Hayward et al., 2005)
      N - Atg8a
      proximity ligation assay, fluorescence microscopy
      (Katz et al., 2025)
      N - baz
      anti tag coimmunoprecipitation, anti tag western blot
      (Wu et al., 2024)
      N - Caf1-55
      anti tag coimmunoprecipitation, anti tag western blot
      (Yu et al., 2013)
      N - Caf1-105
      anti tag coimmunoprecipitation, anti tag western blot
      (Yu et al., 2013)
      N - Caf1-180
      anti tag coimmunoprecipitation, anti tag western blot
      (Yu et al., 2013)
      N - Cdk8
      pull down, western blot
      (Park et al., 2003)
      N - CG10916
      experimental knowledge based
      (Guruharsha et al., 2011)
      N - Chi
      anti tag coimmunoprecipitation, western blot, pull down
      (Sachan et al., 2015)
      N - cno
      anti tag coimmunoprecipitation, western blot, autoradiography, anti bait coimmunoprecipitation, two hybrid
      (Pérez-Gómez et al., 2013, Carmena et al., 2006)
      N - crb
      proximity ligation assay, fluorescence microscopy
      (Nemetschke and Knust, 2016)
      N - D
      affinity technology, tag visualisation by alkaline phosphatase activity
      (Pérez et al., 2005)
      N - Dab
      pull down, autoradiography, western blot, anti bait coimmunoprecipitation, anti tag coimmunoprecipitation
      (Kannan et al., 2018, Le Gall et al., 2008, Le Gall and Giniger, 2004, Giniger, 1998)
      N - Delta
      affinity technology, inferred by author, bead aggregation assay, fluorescence microscopy, enzyme linked immunosorbent assay, anti bait coimmunoprecipitation, western blot, tag visualisation, phenotype-based detection assay, cross-linking study, pull down, tag visualisation by alkaline phosphatase activity, atomic force microscopy
      (Pandey et al., 2019, Yamamoto et al., 2012, Acar et al., 2008, Pei and Baker, 2008, Xu et al., 2007, Parks et al., 2006, Ahimou et al., 2004, Bruckner et al., 2000, Wesley and Saez, 2000, Wesley, 1999, Rebay et al., 1991, Fehon et al., 1990)
      N - dsh
      pull down, western blot, autoradiography, anti bait coimmunoprecipitation, two hybrid
      (Capilla et al., 2012, Muñoz-Descalzo et al., 2010, Hayward et al., 2005, Ramain et al., 2001, Axelrod et al., 1996)
      N - dx
      anti tag coimmunoprecipitation, western blot, nuclear magnetic resonance, two hybrid, cosedimentation in solution, predetermined participant, anti tag western blot
      (Carter et al., 2023, Shimizu et al., 2014, Muñoz-Descalzo et al., 2010, Matsuno et al., 1995, Diederich et al., 1994)
      N - Eip75B
      anti tag coimmunoprecipitation, western blot, proximity ligation assay, fluorescence microscopy
      (Wang et al., 2024)
      N - Eogt
      enzymatic study
      (Sakaidani et al., 2011)
      N - htk
      anti tag coimmunoprecipitation, western blot, anti bait coimmunoprecipitation, two hybrid, anti tag western blot
      (Singh et al., 2019)
      N - Kap-α3
      two hybrid, pull down, western blot, anti tag coimmunoprecipitation
      (Sachan et al., 2013)
      N - kuz
      protease assay, western blot, inferred by author
      (Pan and Rubin, 1997)
      N - lilli
      anti tag coimmunoprecipitation, anti tag western blot
      (Liu et al., 2017)
      N - mam
      three hybrid, protein three hybrid, anti tag coimmunoprecipitation, peptide massfingerprinting
      (Gahr et al., 2019, Nagel et al., 2017, Yuan et al., 2016, Maier et al., 2011, Veraksa et al., 2005, Petcherski and Kimble, 2000)
      N - mask
      anti bait coimmunoprecipitation, western blot, two hybrid
      (Sarkar et al., 2025)
      N - MED17
      pull down, western blot
      (Park et al., 2003)
      N - MED21
      pull down, western blot
      (Park et al., 2003)
      N - MED31
      pull down, western blot
      (Park et al., 2003)
      N - msn
      coimmunoprecipitation, western blot
      (Mishra et al., 2015)
      N - ND-30
      anti tag coimmunoprecipitation, western blot
      (Lee et al., 2013)
      N - Nedd4
      pull down, western blot, anti tag coimmunoprecipitation
      (Sakata et al., 2004)
      N - numb
      two hybrid, pull down, autoradiography
      (Guo et al., 1996)
      N - Ofut1
      anti tag coimmunoprecipitation, western blot, anti tag western blot
      (Sasamura et al., 2007, Okajima et al., 2005)
      N - Pink1
      anti tag coimmunoprecipitation, western blot
      (Lee et al., 2013)
      N - Pngl
      experimental knowledge based
      (Guruharsha et al., 2011)
      N - polo
      experimental knowledge based
      (Guruharsha et al., 2011)
      N - pyd
      anti tag coimmunoprecipitation, anti tag western blot
      (Shimizu et al., 2017)
      N - Rbfox1
      anti bait coimmunoprecipitation, western blot
      (Shukla et al., 2017)
      N - sca
      anti tag coimmunoprecipitation, western blot
      (Petruccelli et al., 2018)
      N - Ser
      inferred by author, anti bait coimmunoprecipitation, cross-linking study, western blot, pull down, tag visualisation by alkaline phosphatase activity, affinity technology, fluorescence microscopy, confocal microscopy, bead aggregation assay, tag visualisation, anti tag coimmunoprecipitation
      (Pandey et al., 2019, Fleming et al., 2013, Pérez-Gómez et al., 2013, Yamamoto et al., 2012, Kim et al., 2010, Xu et al., 2007, Pérez et al., 2005, Wesley, 1999, Rebay et al., 1991)
      N - shg
      anti bait coimmunoprecipitation, western blot
      (Sasaki et al., 2007)
      N - spdo
      anti bait coimmunoprecipitation, western blot, anti tag coimmunoprecipitation, anti tag western blot, pull down
      (Upadhyay et al., 2013, O'Connor-Giles and Skeath, 2003)
      N - Su(dx)
      pull down, molecular weight estimation by staining, isothermal titration calorimetry, predetermined participant, autoradiography, nuclear magnetic resonance, western blot
      (Yao et al., 2018, Jennings et al., 2007, Wilkin et al., 2004)
      N - Su(H)
      protein three hybrid, isothermal titration calorimetry, predetermined participant, two hybrid, pull down, autoradiography, anti tag coimmunoprecipitation, peptide massfingerprinting, anti bait coimmunoprecipitation, western blot, cosedimentation, molecular sieving, molecular weight estimation by staining, anti tag western blot, electrophoretic mobility shift assay, electrophoretic mobility supershift assay, coimmunoprecipitation, electron microscopy, three hybrid
      (Gagliani et al., 2022, Koca et al., 2022, Gahr et al., 2019, Gomez-Lamarca et al., 2018, Nagel et al., 2017, Xu et al., 2017, Yuan et al., 2016, Auer et al., 2015, Contreras et al., 2015, Brockmann et al., 2014, Mishra et al., 2014, Sachan et al., 2013, Kurth et al., 2011, Maier et al., 2011, Kelly et al., 2007, Krejci and Bray, 2007, Cave et al., 2005, Veraksa et al., 2005, Le Gall and Giniger, 2004, Kitagawa et al., 2001, Petcherski and Kimble, 2000, Kidd et al., 1998, Guo et al., 1996, Fortini and Artavanis-Tsakonas, 1994)
      N - Taf9
      anti tag coimmunoprecipitation, anti tag western blot
      (Xie et al., 2014)
      N - Traf6
      two hybrid, anti tag coimmunoprecipitation, western blot, pull down
      (Mishra et al., 2014)
      N - trio
      anti bait coimmunoprecipitation, western blot
      (Kannan et al., 2018, Le Gall et al., 2008)
      N - uif
      anti tag coimmunoprecipitation, anti tag western blot
      (Loubéry et al., 2014)
      N - Vps26
      anti tag coimmunoprecipitation, anti tag western blot
      (Li et al., 2018)
      N - Vps35
      anti tag coimmunoprecipitation, anti tag western blot
      (Li et al., 2018)
      N - wg
      anti bait coimmunoprecipitation, cross-linking study, western blot
      (Wesley, 1999)
      N - wry
      inferred by author, confocal microscopy, bead aggregation assay
      (Kim et al., 2010)
      N - yki
      proximity ligation assay, fluorescence microscopy
      (Wang et al., 2024)
      N - zip
      anti bait coimmunoprecipitation, western blot, Identification by mass spectrometry, anti tag western blot
      (Verma et al., 2024)
      RNA-protein
      Interacting group
      Assay
      References
      N - Sxl
      nucleic acid uv cross-linking assay, autoradiography, anti bait coimmunoprecipitation, quantitative reverse transcription pcr
      (Penn and Schedl, 2007)
      N - TBPH
      pull down, quantitative reverse transcription pcr
      (Swain et al., 2016)
      Delta
      RNA-RNA
      Interacting group
      Assay
      References
      Delta - mir-1
      luminiscence technology, immunohistochemistry
      (Kwon et al., 2005)
      protein-protein
      Interacting group
      Assay
      References
      Delta
      atomic force microscopy, bead aggregation assay, fluorescence microscopy, inferred by author
      (Parks et al., 2006, Ahimou et al., 2004)
      Delta - kuz
      protease assay, western blot, inferred by author
      (Qi et al., 1999)
      Delta - mib1
      enzymatic study, anti tag coimmunoprecipitation, anti tag western blot
      (Daskalaki et al., 2011, Lai et al., 2005)
      Delta - N
      affinity technology, inferred by author, bead aggregation assay, fluorescence microscopy, enzyme linked immunosorbent assay, anti bait coimmunoprecipitation, western blot, tag visualisation, phenotype-based detection assay, cross-linking study, pull down, tag visualisation by alkaline phosphatase activity, atomic force microscopy
      (Pandey et al., 2019, Yamamoto et al., 2012, Acar et al., 2008, Pei and Baker, 2008, Xu et al., 2007, Parks et al., 2006, Ahimou et al., 2004, Bruckner et al., 2000, Wesley and Saez, 2000, Wesley, 1999, Rebay et al., 1991, Fehon et al., 1990)
      Delta - neur
      anti tag coimmunoprecipitation, anti tag western blot, pull down, western blot, anti bait coimmunoprecipitation, enzymatic study
      (Liu et al., 2012, Daskalaki et al., 2011, Fontana and Posakony, 2009, Commisso and Boulianne, 2007, Skwarek et al., 2007, Pitsouli and Delidakis, 2005)
      Delta - Ofut1
      anti tag coimmunoprecipitation, anti tag western blot
      (Okajima et al., 2005)
      Delta - sca
      anti bait coimmunoprecipitation, western blot, anti tag coimmunoprecipitation, coimmunoprecipitation, affinity technology, fluorescence microscopy
      (Petruccelli et al., 2018, Mok et al., 2005)
      Ser
      RNA-RNA
      Interacting group
      Assay
      References
      Ser - mir-8
      luminiscence technology, necessary binding region, immunohistochemistry, fluorescence technology
      (Vallejo et al., 2011)
      protein-protein
      Interacting group
      Assay
      References
      Ser - cno
      anti tag coimmunoprecipitation, western blot
      (Pérez-Gómez et al., 2013)
      Ser - mib1
      anti tag coimmunoprecipitation, anti tag western blot
      (Lai et al., 2005)
      Ser - N
      inferred by author, anti bait coimmunoprecipitation, cross-linking study, western blot, pull down, tag visualisation by alkaline phosphatase activity, affinity technology, fluorescence microscopy, confocal microscopy, bead aggregation assay, tag visualisation, anti tag coimmunoprecipitation
      (Pandey et al., 2019, Fleming et al., 2013, Pérez-Gómez et al., 2013, Yamamoto et al., 2012, Kim et al., 2010, Xu et al., 2007, Pérez et al., 2005, Wesley, 1999, Rebay et al., 1991)
      Ser - neur
      anti bait coimmunoprecipitation, western blot, anti tag coimmunoprecipitation, anti tag western blot
      (Liu et al., 2012, Glittenberg et al., 2006, Pitsouli and Delidakis, 2005)
      Alleles Reported to Model Human Disease (Disease Ontology) (38 alleles)
      N
      Models Based on Experimental Evidence ( 13 )
      Allele
      Disease
      Evidence
      References
      NDN.UAS.cUa
      model of  cancer
      CEA with Ras85DV12.cUa.UAS
      (Lee et al., 2021)
      CEA
      (Lee et al., 2021, Tamamouna et al., 2021)
      CEA with Ras85DV12.UAS
      (Xu et al., 2023)
      model of  intestinal cancer
      CEA
      (Singh et al., 2016)
      NICN.UAS
      model of  cancer
      CEA
      (Wang et al., 2021, Harsh and Eleftherianos, 2020)
      CEA with Mef2UAS.cUa
      (Harsh and Eleftherianos, 2020)
      model of  carcinoma
      CEA with Mef2UAS.cUa
      (Pallavi et al., 2012)
      CEA with egrUAS.cUa
      (Pallavi et al., 2012)
      model of  brain cancer
      CEA
      (Wang et al., 2006)
      NUAS.cMa
      model of  adenoid cystic carcinoma
      CEA with Hsap\MYBΔC.UAS, XpdRNAi.UAS.012.1, ftRNAi.UAS.012.1, kugRNAi.UAS.012.1
      (Bangi et al., 2021)
      CEA with Hsap\MYBΔC.UAS, XpdRNAi.UAS.012.2, ftRNAi.UAS.012.2, kugRNAi.UAS.012.2
      (Bangi et al., 2021)
      NUAS.icd
      model of  central nervous system cancer
      CEA
      (Zacharioudaki et al., 2016)
      NRNAi.P.UAS
      model of  intestinal cancer
      CEA
      (Chen et al., 2021)
      CEA with mysGD15002
      (Chen et al., 2021)
      model of  cancer
      CEA with Ras85DV12.UAS
      (Tamamouna et al., 2021)
      Nintra.GS.UAS
      model of  neurodegenerative disease
      CEA
      (Loewen et al., 2018)
      NUAS.Tag:V5,Tag:polyHis
      model of  brain cancer
      CEA
      (Ojha et al., 2022)
      model of  cancer
      CEA
      (Khaket et al., 2024)
      NGD144
      model of  alcohol use disorder
      CEA
      (Petruccelli et al., 2018)
      NGD14477
      model of  intestinal cancer
      CEA
      (Banreti et al., 2022)
      model of  colorectal cancer
      CEA
      (Zipper et al., 2022)
      model of  cancer
      CEA
      (Brás et al., 2021)
      NHD_CFD01184
      model of  intestinal cancer
      CEC with Spyo\Cas9UAS.uM.-FRT
      (Bahuguna et al., 2021)
      CEC with Spyo\Cas9UAS.P2
      (Bahuguna et al., 2021)
      NCA.UAS.cUa
      model of  central nervous system cancer
      CEA
      (Magadi et al., 2020, Zacharioudaki et al., 2016)
      model of  carcinoma
      CEA with l(2)glGD4047
      (Jevitt et al., 2021)
      model of  adenocarcinoma
      CEA
      (Casas-Tintó et al., 2016)
      CEA with wupAf06492
      (Casas-Tintó et al., 2016)
      model of  cervical cancer
      CEA with Hsap\UBE3AUAS.cRa, HPV18\E6UAS.Tag:MYC
      (Padash Barmchi et al., 2016)
      NRNAi.UAS.cUa
      model of  intestinal cancer
      CEA
      (Ayyaz et al., 2015)
      NΔE.UAS
      model of  central nervous system cancer
      CEA
      (Voutyraki et al., 2023, Voutyraki et al., 2022)
      Modifiers Based on Experimental Evidence ( 22 )
      Allele
      Disease
      Interaction
      References
      NDN.UAS.cUa
      model of  cancer
      is ameliorated by wgJF01257
      (Lee et al., 2021)
      is ameliorated by Atg1UAS.cSa
      (Lee et al., 2021)
      ameliorates  carcinoma
      modeled by Vps25N55
      (Herz et al., 2006)
      modeled by ph-d505, ph-p505
      (Beira et al., 2018)
      ameliorates  intestinal cancer
      modeled by ykiS111A.S168A.S250A.UAS.Tag:V5
      (Pranoto et al., 2023)
      NECN.UAS
      ameliorates  cancer
      modeled by Src64BUAS.cCa
      (Enomoto et al., 2021)
      NFLN.ΔCdc10.UAS
      ameliorates  C9orf72 frontotemporal dementia and/or amyotrophic lateral sclerosis
      modeled by Zzzz\poly-GR80.UAS.Tag:FLAG
      (Li et al., 2020)
      NICN.UAS
      model of  cancer
      is ameliorated by CycEHMS00060
      (Wang et al., 2021)
      is ameliorated by DrefJF02232
      (Wang et al., 2021)
      is ameliorated by RPA1GL00350
      (Wang et al., 2021)
      is ameliorated by SMC5HMC04343
      (Wang et al., 2021)
      is ameliorated by fzrGD9960
      (Wang et al., 2021)
      is ameliorated by mei-41GL00284
      (Wang et al., 2021)
      is ameliorated by mre11HMC02995
      (Wang et al., 2021)
      is exacerbated by stgUAS.Tag:HA
      (Wang et al., 2021)
      is ameliorated by stgRNAi.UAS.cUa
      (Wang et al., 2021)
      exacerbates  cancer
      modeled by Src64BUAS.cCa
      (Enomoto et al., 2021)
      modeled by HipkUAS.Tag:HA
      (Blaquiere et al., 2018)
      exacerbates  carcinoma
      modeled by scrib1
      (Brumby and Richardson, 2003)
      NUAS.icd
      model of  central nervous system cancer
      is ameliorated by hthC1
      (Zacharioudaki et al., 2016)
      is exacerbated by miraHMS00347
      (Zacharioudaki et al., 2016)
      is ameliorated by svp1
      (Zacharioudaki et al., 2016)
      is ameliorated by hthB2
      (Zacharioudaki et al., 2016)
      NRNAi.P.UAS
      model of  intestinal cancer
      is exacerbated by CatHMS00990
      (Chen et al., 2021)
      is ameliorated by CatUAS.cUa
      (Chen et al., 2021)
      is ameliorated by Sod2UAS.cMa
      (Chen et al., 2021)
      is exacerbated by cncHMS00650
      (Chen et al., 2021)
      is exacerbated by Sod2HMS00499
      (Chen et al., 2021)
      is ameliorated by cncUAS.cSa
      (Chen et al., 2021)
      ameliorates  carcinoma
      modeled by Vps25A3
      (Vaccari and Bilder, 2005)
      Nfa-swb
      ameliorates  Parkinson's disease 1
      modeled by Hsap\SNCAUAS.cJa
      (Rousseaux et al., 2018)
      Nintra.GS.UAS
      ameliorates  cancer
      modeled by Pvrλ.UASp.Tag:MYC
      (Wang et al., 2023)
      exacerbates  carcinoma
      modeled by NkapGD11807
      (Guo et al., 2024)
      Nl1N-ts1
      ameliorates  brain cancer
      modeled by aurA8839
      (Wang et al., 2006)
      ameliorates  frontotemporal dementia and/or amyotrophic lateral sclerosis 7
      modeled by Hsap\CHMP2BIntron5.UAS
      (Wilson et al., 2020)
      Nnd-1
      ameliorates  Schimke immuno-osseous dysplasia
      modeled by Marcal1UAS.cBa
      (Morimoto et al., 2016)
      Nnd-3
      ameliorates  Schimke immuno-osseous dysplasia
      modeled by Marcal1UAS.cBa
      (Morimoto et al., 2016)
      NUAS.Tag:V5,Tag:polyHis
      model of  brain cancer
      is ameliorated by ND-49HM05059
      (Ojha et al., 2022)
      is ameliorated by Scer\NDI1UAS.cBa
      (Ojha et al., 2022)
      ameliorates  C9orf72 frontotemporal dementia and/or amyotrophic lateral sclerosis
      modeled by Zzzz\poly-GR80.UAS.Tag:FLAG
      (Li et al., 2020)
      modeled by Zzzz\poly-GA80.UAS.Tag:FLAG
      (Li et al., 2020)
      modeled by Zzzz\poly-PR80.UAS.Tag:FLAG
      (Li et al., 2020)
      model of  cancer
      is ameliorated by TER94JF03402
      (Khaket et al., 2024)
      is ameliorated by peloKK101657
      (Khaket et al., 2024)
      is ameliorated by peloUAS.ORF.GW.Tag:HA
      (Khaket et al., 2024)
      is ameliorated by pixJF01190
      (Khaket et al., 2024)
      is ameliorated by pixUAS.ORF.Tag:HA
      (Khaket et al., 2024)
      is ameliorated by CG11414HMJ23120
      (Khaket et al., 2024)
      is ameliorated by Cnot4JF03203
      (Khaket et al., 2024)
      is ameliorated by SARS-CoV-2\nsp1UAS.cZa
      (Khaket et al., 2024)
      NJF01637
      ameliorates  carcinoma
      modeled by l(2)glGD4047
      (Chatterjee et al., 2023)
      modeled by NkapGD11807
      (Guo et al., 2024)
      NGD14477
      model of  intestinal cancer
      is exacerbated by PcmtGD8637
      (Banreti et al., 2022)
      is exacerbated by PcmtRNAi.UAS
      (Banreti et al., 2022)
      ameliorates  cancer
      modeled by Src64BUAS.cCa
      (Enomoto et al., 2021)
      model of  colorectal cancer
      is ameliorated by mir-277UAS.cSa
      (Zipper et al., 2022)
      ameliorates  brain cancer
      modeled by Vps35E42
      (Li et al., 2018)
      exacerbates  cancer
      modeled by Mps1GL00184
      (Brás et al., 2021)
      modeled by Mps1HMS01546
      (Brás et al., 2021)
      modeled by mad2GLC01381
      (Brás et al., 2021)
      model of  cancer
      is exacerbated by Mps1GL00184
      (Brás et al., 2021)
      is exacerbated by Mps1HMS01546
      (Brás et al., 2021)
      is exacerbated by mad2GLC01381
      (Brás et al., 2021)
      NNICD.UASp
      exacerbates  carcinoma
      modeled by l(2)glGD4047
      (Chatterjee et al., 2023)
      NsfGFP,Tag:FLAG
      ameliorates  Adams-Oliver syndrome
      modeled by Su(H)T4
      (Gagliani et al., 2022)
      NJF02959
      ameliorates  rhabdoid cancer
      modeled by Snr1HMS00363, BacA\p35UAS.cHa
      (Xie et al., 2017)
      NRNAi.lexAop
      ameliorates  carcinoma
      modeled by EgfrUAS.cBa, psqKK111691
      (Boukhatmi et al., 2020)
      NUAS.cGa
      ameliorates  autism spectrum disorder
      modeled by Nrx-IV4304
      (Grice et al., 2015)
      NCA.UAS.cUa
      model of  central nervous system cancer
      is ameliorated by zfh1UAS.cPa
      (Magadi et al., 2020)
      is ameliorated by gcmUAS.ORF.GW.Tag:HA
      (Magadi et al., 2020)
      exacerbates  carcinoma
      modeled by l(2)glGD4047
      (Jevitt et al., 2021)
      NRNAi.UAS.cUa
      model of  intestinal cancer
      is ameliorated by SmoxJF02320
      (Ayyaz et al., 2015)
      NΔE.UAS
      model of  central nervous system cancer
      is exacerbated by InRA1325D.UAS
      (Voutyraki et al., 2023)
      is ameliorated by InRK1409A.UAS
      (Voutyraki et al., 2023)
      is ameliorated by ImpHMS01168
      (Voutyraki et al., 2023)
      Delta
      Models Based on Experimental Evidence ( 2 )
      Modifiers Based on Experimental Evidence ( 9 )
      Allele
      Disease
      Interaction
      References
      DeltaHMS01309
      ameliorates  brain cancer
      modeled by Vps35E42
      (Li et al., 2018)
      Delta9P
      ameliorates  multiple endocrine neoplasia type 2B
      modeled by RetMEN2B.GMR
      (Read et al., 2005)
      ameliorates  multiple endocrine neoplasia type 2A
      modeled by RetMEN2A.GMR
      (Read et al., 2005)
      DeltaRevF10
      ameliorates  cancer
      modeled by Src64BUAS.cCa
      (Enomoto et al., 2021)
      ameliorates  amyotrophic lateral sclerosis
      modeled by Hsap\TARDBPUAS.cHa, SerRX82
      (Zhan et al., 2013)
      DeltaS049520
      ameliorates  multiple endocrine neoplasia type 2B
      modeled by RetMEN2B.GMR
      (Read et al., 2005)
      ameliorates  multiple endocrine neoplasia type 2A
      modeled by RetMEN2A.GMR
      (Read et al., 2005)
      DeltaUAS.cDa
      model of  cancer
      is exacerbated by Not3KG10496
      (De Keersmaecker et al., 2013)
      is exacerbated by Not3GD4068
      (De Keersmaecker et al., 2013)
      is exacerbated by Not3KK102144
      (De Keersmaecker et al., 2013)
      is exacerbated by ItgaPS4KK105806
      (Zhai et al., 2012)
      is exacerbated by ctRNAi.cGa.UAS
      (Zhai et al., 2012)
      is exacerbated by mysHMS00043
      (Zhai et al., 2012)
      DeltaJF02867
      ameliorates  carcinoma
      modeled by EgfrUAS.cBa, psqKK111691
      (Boukhatmi et al., 2020)
      Delta05151
      exacerbates  ataxia telangiectasia
      modeled by tefuRNAi.UAS
      (Rimkus et al., 2008)
      DeltaKK107312
      exacerbates  Alzheimer's disease
      modeled by PsnRNAi.UAS.3
      (Kang et al., 2024)
      DeltaDN.UAS
      ameliorates  brain cancer
      modeled by Vps35E42
      (Li et al., 2018)
      DOES NOT ameliorate  cancer
      modeled by HipkUAS.Tag:HA
      (Blaquiere et al., 2018)
      ameliorates  cancer
      modeled by Src64BUAS.cCa
      (Enomoto et al., 2021)
      Ser
      Models Based on Experimental Evidence ( 2 )
      Allele
      Disease
      Evidence
      References
      SerDN.UAS
      model of  dilated cardiomyopathy
      CEA
      (Kim et al., 2010)
      SerRNAi.UAS.cUa
      model of  dilated cardiomyopathy
      CEA
      (Kim et al., 2010)
      Modifiers Based on Experimental Evidence ( 2 )
      Alleles Representing Disease-Implicated Variants
      Genetic Tools, Stocks and Reagents
      Sources of Stocks
      Contact lab of origin for a reagent not available from a public stock center.
      Bloomington Stock Center Disease Page
      Related mammalian, viral, bacterial, or synthetic transgenes
      Allele
      Transgene
      Publicly Available Stocks
      Delta::Hsap\JAG1UAS.cytoWT
      M{UAS-Delta::J1.cytoWT}
      Delta::Hsap\JAG1N-ala.C-ala.UAS
      M{UAS-Delta::J1.N-ala.C-ala}
      Delta::Hsap\JAG1KR.UAS
      M{UAS-Delta::J1.KR}
      Selected Drosophila transgenes
      Allele
      Transgene
      Publicly Available Stocks
      NHD_CFD01184
      P{HD_CFD01184}
      P{hsFLP}1, y1 w1118; P{HD_CFD01184}attP40/CyO-GFP
      NUAS.cGa
      P{UAS-N.G}
      NCA.UAS.cUa
      P{UAS-N.CA.U}
      NΔE.UAS
      P{UAS-Delta.DeltaE}
      y1 w*; P{UAS-Delta.ΔE}B2a2
      NECN.UAS
      P{UAS-N.ECN}
      P{hsFLP}12, y1 w*; P{UAS-N.ECN}2; MKRS/TM2
      NICN.UAS
      P{UAS-N.ICN}
      NFLN.ΔCdc10.UAS
      P{UAS-N.FLNDeltaCdc10}
      Nintra.GS.UAS
      P{UAS-N.intra.GS}
      NUAS.icd
      P{UAS-N.icd}
      NDN.UAS.cUa
      P{UAS-N.DN.U}
      NUAS.cMa
      P{UAS-Nfull}
      w*; P{UAS-Nfull}3
      NUAS.Tag:V5,Tag:polyHis
      P{UAS-N.V5}
      NsfGFP,Tag:FLAG
      PBac{N-GFP.FLAG}
      y1 w*; PBac{N-GFP.FLAG}VK00033/TM6C, Sb1
      NNICD.UASp
      P{UASp-N.NICD}
      DeltaDN.UAS
      P{UAS-Delta.DN}
      w1118; Dr1 e1/TM3, P{UAS-Delta.DN}TJ1, Sb1
      DeltaUAS.cDa
      P{UAS-Delta.D}
      SerDN.UAS
      P{UAS-Ser.DN}
      w*; P{UAS-Ser.DN}3
      RNAi constructs available
      Allele
      Transgene
      Publicly Available Stocks
      NRNAi.lexAop
      P{lexAop-N.RNAi}
      NRNAi.13xlexAop.cZa
      P{13xlexAop-N.RNAi.Z}
      NlexAop.HMS00009
      M{lexAop-HMS00009}
      NRNAi.UAS.cUa
      P{UAS-N.RNAi.U}
      NRNAi.P.UAS
      P{UAS-N.RNAi.M}
      w*; P{UAS-N.RNAi.P}9G
      NRNAi.G.UAS
      P{UAS-N.RNAi.G}
      NGD144
      P{GD144}
      w1118; P{GD144}v1112
      NGD14477
      P{GD14477}
      w1118; P{GD14477}v27228/TM3
      NGD2640
      P{GD2640}
      NKK102890
      P{KK102890}
      P{KK102890}VIE-260B
      NJF02959
      P{TRiP.JF02959}
      y1 v1; P{TRiP.JF02959}attP2
      NJF01637
      P{TRiP.JF01637}
      y1 v1; P{TRiP.JF01637}attP2
      NJF01043
      P{TRiP.JF01043}
      y1 v1; P{TRiP.JF01043}attP2
      NJF01053
      P{TRiP.JF01053}
      y1 v1; P{TRiP.JF01053}attP2
      NJF01356
      P{TRiP.JF01356}
      y1 v1; P{TRiP.JF01356}attP2
      NJF01693
      P{TRiP.JF01693}
      y1 v1; P{TRiP.JF01693}attP2
      NJF01047
      P{TRiP.JF01047}
      NJF01892
      P{TRiP.JF01892}
      NJF01891
      P{TRiP.JF01891}
      NJF01893
      P{TRiP.JF01893}
      NJF01358
      P{TRiP.JF01358}
      NJF02179
      P{TRiP.JF02179}
      NNIG.3936R
      P{NIG.3936R}
      P{NIG.3936R}2
      NHMS00001
      P{TRiP.HMS00001}
      y1 v1; P{TRiP.HMS00001}attP2
      NHMS00009
      P{TRiP.HMS00009}
      y1 v1; P{TRiP.HMS00009}attP2
      NRNAi.UAS
      P{UAS-N.RNAi}
      NGL00092
      P{TRiP.GL00092}
      y1 sc* v1 sev21; P{TRiP.GL00092}attP2
      NGLV21004
      P{TRiP.GLV21004}
      y1 sc* v1 sev21; P{TRiP.GLV21004}attP2
      NRNAi.UAS.cHa
      P{UAS-N.RNAi.H}
      NTRiP.valium1
      P{TRiP.N.valium1}
      DeltaRNAi.UAS.XK8
      P{UAS-Delta.RNAi.XK8}
      DeltaRNAi.UAS.cCa
      P{UAS-Delta.RNAi.C}
      DeltaRNAi.UAS.cUa
      P{UAS-Delta.RNAi.U}
      DeltaGD146
      P{GD146}
      w1118; P{GD146}v3720
      DeltaGD2642
      P{GD2642}
      w1118; P{GD2642}v37287
      DeltaGD14459
      P{GD14459}
      DeltaJF02867
      P{TRiP.JF02867}
      y1 v1; P{TRiP.JF02867}attP2
      DeltaHMS01309
      P{TRiP.HMS01309}
      y1 sc* v1 sev21; P{TRiP.HMS01309}attP2
      DeltaKK107312
      P{KK107312}
      P{KK107312}VIE-260B
      DeltaGL00520
      P{TRiP.GL00520}
      y1 sc* v1 sev21; P{TRiP.GL00520}attP40
      SerGD147
      P{GD147}
      SerGD14453
      P{GD14453}
      w1118; P{GD14453}v27172
      SerGD2643
      P{GD2643}
      SerKK108416
      P{KK108416}
      P{KK108416}VIE-260B
      SerJF03140
      P{TRiP.JF03140}
      y1 v1; P{TRiP.JF03140}attP2
      SerRNAi.UAS.cUa
      P{UAS-Ser.RNAi.U}
      SerHMS01179
      P{TRiP.HMS01179}
      y1 sc* v1 sev21; P{TRiP.HMS01179}attP2
      Selected Drosophila classical alleles
      Allele
      Allele class
      Mutagen
      Publicly Available Stocks
      NETX52
      amorphic allele - genetic evidence
      Delta2-3 transposase
      N55e11
      loss of function allele
      spontaneous
      N55e11 P{neoFRT}19A/FM7c
      Nfa-g
      loss of function allele
      spontaneous
      wa Nfa-g; Rho1E3.10/CyO
      N264-40
      amorphic allele - genetic evidence
      X ray
      wa N264-40 rb1/C(1)DX, y1 f1; Dp(1;2)72c21/+
      NFE1
      loss of function allele
      X ray
      Nl1N-ts1
      ethyl methanesulfonate
      y1 Nl1N-ts1 g2 f1/C(1)DX, y1 w1 f1
      NS-317
      loss of function allele
      ethyl methanesulfonate
      Nl1N-2
      amorphic allele - genetic evidence
      gamma ray
      NFX9
      loss of function allele
      X ray
      NFE2
      loss of function allele
      X ray
      NFX8
      loss of function allele
      X ray
      N1
      loss of function allele
      y1 w* N1/FM7c, P{GAL4-twi.G}108.4, P{UAS-2xEGFP}AX
      NFX4
      loss of function allele
      X ray
      NeU10
      loss of function allele
      ethyl methanesulfonate
      NeAS7b
      loss of function allele
      ethyl methanesulfonate
      N60g11
      amorphic allele - genetic evidence
      X ray
      NFX2
      loss of function allele
      X ray
      NXL86
      loss of function allele
      ethyl methanesulfonate
      N2
      loss of function allele
      NFX5
      loss of function allele
      X ray
      NFX7
      loss of function allele
      X ray
      NYi66
      loss of function allele
      ethyl methanesulfonate
      N264-47
      loss of function allele
      X ray
      N264-39
      loss of function allele
      X ray
      wch2 N264-39/FM4, Bar+
      NX81K1
      loss of function allele
      Nfa-swb
      X ray
      wa Nfa-swb
      N3
      loss of function allele
      N3b22
      loss of function allele
      ethyl methanesulfonate
      NFX6
      loss of function allele
      X ray
      NXK11
      amorphic allele - genetic evidence
      Nnd-1
      loss of function allele
      spontaneous
      Nnd-1 rb1
      NM1
      loss of function allele
      ethyl methanesulfonate
      NFX3
      loss of function allele
      X ray
      NFX1
      loss of function allele
      X ray
      NX114
      loss of function allele
      ethyl methanesulfonate
      N66l2
      amorphic allele - genetic evidence
      Nnd-3
      X ray
      Nnd-3 rb1/C(1)DX, y1 f1
      Delta05151
      P-element activity
      DeltaS049520
      Delta2-3 transposase
      DeltaΔExon2
      amorphic allele - molecular evidence
      CRISPR/Cas9
      DeltaBX44
      amorphic allele - genetic evidence
      X ray
      DeltaFE27
      amorphic allele - genetic evidence
      ethyl methanesulfonate
      Delta9P
      amorphic allele - genetic evidence
      ethyl methanesulfonate
      ru1 hry1 Diap1th-1 st1 cu1 sr1 Delta9P es ca1/TM3, Sb1
      DeltaBE30
      amorphic allele - genetic evidence
      ethyl methanesulfonate
      DeltaB2
      amorphic allele - genetic evidence
      ethyl methanesulfonate
      DeltaB2 e1/TM6C, Sb1
      Delta3
      loss of function allele
      spontaneous
      Delta3/In(3R)C, sprd1 e1
      DeltaRevF10
      loss of function allele
      DeltaRevF10 e* SerRX82 P{neoFRT}82B/TM6B, Tb1
      DeltaM1
      loss of function allele
      X ray
      DeltaBE32
      amorphic allele - genetic evidence
      ethyl methanesulfonate
      DeltaM
      loss of function allele
      spontaneous
      DeltaX
      amorphic allele - genetic evidence
      DeltaX/In(3L)P, In(3R)P
      Deltavi3
      loss of function allele
      ethyl methanesulfonate
      Deltavi1
      loss of function allele
      ethyl methanesulfonate
      DeltaCE21
      amorphic allele - genetic evidence
      ethyl methanesulfonate
      DeltaBX38
      amorphic allele - genetic evidence
      X ray
      DeltaCE23
      amorphic allele - genetic evidence
      ethyl methanesulfonate
      DeltaBX39
      amorphic allele - genetic evidence
      X ray
      DeltaKX1
      amorphic allele - genetic evidence
      X ray
      DeltaM3
      loss of function allele
      ethyl methanesulfonate
      DeltaM2
      loss of function allele
      spontaneous
      DeltaEP3816.A
      loss of function allele
      Delta2-3 transposase
      Ser1
      spontaneous
      P{NIG.11658R}3/TM3, P{ActGFP}JMR2, Ser1
      Sermf553
      loss of function allele
      minos activity
      SerCS94
      amorphic allele - genetic evidence
      SerRX82
      loss of function allele
      X ray
      DeltaRevF10 e* SerRX82 P{neoFRT}82B/TM6B, Tb1
      SerRX107
      loss of function allele
      X ray
      SerRX106
      amorphic allele - genetic evidence
      X ray
      mwh1 red1 e1 SerRX106/TM6B, Tb1
      Serrev2-11
      loss of function allele
      X ray
      References (52)