FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Human Disease Model Report: cancer, epithelial, LLGL-YAP1-related
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General Information
Name
cancer, epithelial, LLGL-YAP1-related
FlyBase ID
FBhh0000590
Disease Ontology Term
Parent Disease
OMIM
Overview

This model of epithelial cancer uses the Drosophila Scribble polarity complex gene l(2)gl in combination with a transcriptional co-activator regulated by the Hippo signaling pathway, the Drosophila gene yki. The highly conserved Scribble polarity complex plays a key role in determining cell polarity and cell proliferation in epithelial cells. One of the human gene orthologous to Dmel\yki, YAP1, is known to play a role in the development and progression of multiple cancers. See also 'cancer, epithelial, Scribble-complex-related' (FBhh0000586) and 'cancer, epithelial, RAS-LLGL-related' (FBhh0000588).

In human, there are two genes orthologous to Dmel\l(2)gl, the cytoskeletal proteins LLGL1 and LLGL2. Classical loss-of-function alleles, RNAi-targeting constructs, and alleles caused by insertional mutagenesis have been generated for l(2)gl; an amorphic allele has been created via gene targeting and recombination. A tagged wild-type transgene of human Hsap\LLGL1 has been introduced into flies; partial heterologous rescue (functional complementation) of the homozygous Dmel\l(2)gl lethal phenotype is observed.

Animals homozygous for loss-of-function mutations of Dmel\l(2)gl typically die during the larval stage. Somatic clones that are homozygous for loss-of-function mutations of l(2)gl exhibit a cell polarity defect; mutant cells are eliminated from mosaic epithelia by cell competition. Multiple physical and many genetic interactions for Dmel\l(2)gl have been described; see below and in the gene report for l(2)gl.

In human, there are two genes orthologous to Dmel\yki, YAP1 and WWTR1. Classical loss-of-function alleles, RNAi-targeting constructs, and alleles caused by insertional mutagenesis have been generated for yki; an amorphic allele has been created via gene targeting and recombination. Multiple UAS constructs of the human Hsap\YAP1 gene have been introduced into flies, including wild-type and genes with mutations eliminating phosphorylation sites.

Animals homozygous for an amorphic mutation of Dmel\yki die as late embryos and early first instar larvae. Somatic clones are difficult to recover and are very small, indicating that complete loss of function is virtually cell-lethal. Models of cancer involving Dmel\yki use a constitutively active form or drive overexpression of a wild-type gene. MANY physical and genetic interactions for Dmel\yki have been described; see below and in the gene report for yki.

Clones of cells with an amorphic mutation of Dmel\l(2)gl and overexpression of Dmel\yki cause overgrowth of mutant cells in the wing pouch, eventually developing into tumors.

[updated Aug. 2017 by FlyBase; FBrf0222196]

Disease Summary Information
Disease Summary: cancer, epithelial, LLGL-YAP1-related
OMIM report
Human gene(s) implicated
Symptoms and phenotype
Genetics
Cellular phenotype and pathology
Molecular information

LLGL1 and LLGL2 encode cortical cytoskeleton proteins found in a complex involved in maintaining cell polarity and epithelial integrity, the Scribble Cell Polarity Complex. This complex plays a role in the initial phase of the establishment of epithelial cell polarity; it is involved in the regulation of mitotic spindle orientation, proliferation, differentiation, and tissue organization of neuroepithelial cells. [Gene Cards, LLGL1, LLGL2; 2017.08.01]

(FlyBase Curators, 2013-)

YAP1 encodes a transcriptional regulator which can act both as a coactivator and a corepressor and is the critical downstream regulatory target in the Hippo signaling pathway; it thus plays a key role development, growth, repair, and homeostasis. YAP1 is known to play a role in the development and progression of multiple cancers. [Gene Cards, YAP1; 2017.08.01]

(FlyBase Curators, 2013-)
External links
Disease synonyms
Ortholog Information
Human gene(s) in FlyBase
Human gene (HGNC)
Symbol / Name
YAP1; Yes1 associated transcriptional regulator
D. melanogaster ortholog (based on DIOPT)
Dmel\yki
(DIOPT, 2013-)
Comments on ortholog(s)

Many to one: 2 human to 1 Drosophila; the second human gene is WWTR1.

Human gene (HGNC)
Symbol / Name
LLGL1; LLGL scribble cell polarity complex component 1
D. melanogaster ortholog (based on DIOPT)
Dmel\l(2)gl
(DIOPT, 2013-)
Comments on ortholog(s)

Many to one: 2 human to 1 Drosophila; the second human gene is LLGL2.

Human gene (HGNC)
Symbol / Name
LLGL2; LLGL scribble cell polarity complex component 2
D. melanogaster ortholog (based on DIOPT)
Dmel\l(2)gl
(DIOPT, 2013-)
Comments on ortholog(s)

Many to one: 2 human to 1 Drosophila; the second human gene is LLGL1.

Other mammalian ortholog(s) used
    D. melanogaster Gene Information (2)
    Dmel\l(2)gl
    Gene Snapshot
    lethal (2) giant larvae (l(2)gl) encodes a tumor suppressor protein that regulates cell polarity and asymmetric cell division. It acts on the basolateral side of epithelial cells, antagonizing the activity of apical complex proteins encoded by baz, par-6 and aPKC. [Date last reviewed: 2019-09-26]
    Molecular function (GO)
    Cellular component (GO)
    Gene Groups / Pathways
    Comments on ortholog(s)

    Moderate- to high-scoring ortholog of human LLGL1 and LLGL2 (1 Drosophila to 2 human). Dmel\l(2)gl shares 34-36% identity and 51-54% 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)
    Dmel\yki
    Gene Snapshot
    yorkie (yki) encodes a transcriptional co-activator protein that is negatively regulated by Hippo signaling, which promotes its cytoplasmic localization. It functions together with DNA binding proteins, such as the product of sd, to activate transcription of a wide range of downstream targets, including genes that promote growth. [Date last reviewed: 2019-03-21]
    Molecular function (GO)
    Cellular component (GO)
    Gene Groups / Pathways
    Comments on ortholog(s)

    Moderate-scoring ortholog of human YAP1 and WWTR1 (1 Drosophila to 2 human). Dmel\yki shares 31% identity and 45% similarity with YAP1; it shares 25% identity and 34% similarity with WWTR1.

    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 (97 groups)
      l(2)gl
      protein-protein
      Interacting group
      Assay
      References
      l(2)gl
      anti tag coimmunoprecipitation, anti tag western blot
      (Betschinger et al., 2005)
      l(2)gl - aPKC
      anti bait coimmunoprecipitation, western blot, experimental knowledge based, proximity ligation assay, fluorescence microscopy, anti tag coimmunoprecipitation, peptide massfingerprinting
      (Portela et al., 2024, Calero-Cuenca et al., 2016, Goh et al., 2013, Guruharsha et al., 2011, Tian and Deng, 2008, Wirtz-Peitz et al., 2008, Betschinger et al., 2003)
      l(2)gl - ball
      anti tag coimmunoprecipitation, western blot
      (Link et al., 2019)
      l(2)gl - ctp
      two hybrid
      (Hu et al., 2017.6.13)
      l(2)gl - dsh
      anti tag coimmunoprecipitation, western blot
      (Kaplan and Tolwinski, 2010)
      l(2)gl - Git
      proximity ligation assay, fluorescence microscopy
      (Portela et al., 2024)
      l(2)gl - mahj
      anti tag coimmunoprecipitation, anti tag western blot
      (Tamori et al., 2010)
      l(2)gl - mts
      anti tag coimmunoprecipitation, western blot
      (Ogawa et al., 2009)
      l(2)gl - Pak
      anti tag coimmunoprecipitation, western blot
      (Bahri et al., 2010)
      l(2)gl - par-1
      anti tag coimmunoprecipitation, western blot, anti bait coimmunoprecipitation
      (Tian and Deng, 2008)
      l(2)gl - par-6
      anti bait coimmunoprecipitation, western blot, anti tag western blot, peptide massfingerprinting, experimental knowledge based, pull down, autoradiography, anti tag coimmunoprecipitation
      (Neumüller et al., 2012, Guruharsha et al., 2011, Wirtz-Peitz et al., 2008, Hutterer et al., 2004, Betschinger et al., 2003)
      l(2)gl - Vap33
      anti tag coimmunoprecipitation, anti tag western blot, proximity ligation assay, fluorescence microscopy, pull down, Identification by mass spectrometry
      (Portela et al., 2024, Portela et al., 2018)
      l(2)gl - Vha44
      anti tag coimmunoprecipitation, anti tag western blot
      (Portela et al., 2018)
      l(2)gl - zip
      anti tag coimmunoprecipitation, anti tag western blot
      (Betschinger et al., 2005)
      yki
      RNA-RNA
      Interacting group
      Assay
      References
      yki - mir-7
      luminiscence technology
      (Yu et al., 2020)
      yki - mir-8
      fluorescence technology
      (Sander et al., 2018)
      RNA-protein
      Interacting group
      Assay
      References
      yki - Rox8
      anti tag coimmunoprecipitation, quantitative reverse transcription pcr
      (Guo et al., 2020)
      protein-protein
      Interacting group
      Assay
      References
      yki - 14-3-3ε
      anti tag coimmunoprecipitation, anti tag western blot, protein complementation assay, peptide massfingerprinting
      (Srivastava et al., 2021, Wang et al., 2016, Hu et al., 2014, Kwon et al., 2013, Wehr et al., 2013, Ren et al., 2010, Oh and Irvine, 2008)
      yki - 14-3-3ζ
      anti tag coimmunoprecipitation, anti tag western blot, peptide massfingerprinting, western blot
      (Srivastava et al., 2021, Wang et al., 2016, Hu et al., 2014, Kwon et al., 2013, Ren et al., 2010, Oh and Irvine, 2008, Dong et al., 2007)
      yki - Ack
      pull down, peptide massfingerprinting, anti tag coimmunoprecipitation, western blot, anti tag western blot
      (Hu et al., 2016, Kwon et al., 2013, Schoenherr et al., 2012)
      yki - Akt
      enzymatic study, western blot
      (Li et al., 2022)
      yki - Atpα
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Kwon et al., 2013)
      yki - Bap111
      anti bait coimmunoprecipitation, peptide massfingerprinting
      (Oh et al., 2013)
      yki - bon
      anti tag coimmunoprecipitation, anti tag western blot, pull down, Identification by mass spectrometry
      (Zhao et al., 2023)
      yki - brm
      anti bait coimmunoprecipitation, peptide massfingerprinting, anti tag coimmunoprecipitation, anti tag western blot
      (Zhu et al., 2015, Jin et al., 2013, Oh et al., 2013)
      yki - CalpA
      anti tag coimmunoprecipitation, Identification by mass spectrometry, anti tag western blot
      (Zhai et al., 2024)
      yki - CalpB
      anti tag coimmunoprecipitation, anti tag western blot
      (Zhai et al., 2024)
      yki - caz
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Kwon et al., 2013)
      yki - cbt
      anti tag coimmunoprecipitation, western blot, anti tag western blot
      (Srivastava et al., 2021, Ruiz-Romero et al., 2015)
      yki - Cdk7
      enzymatic study, western blot
      (Cho et al., 2020)
      yki - CG2186
      anti tag coimmunoprecipitation, anti tag western blot, peptide massfingerprinting
      (Kwon et al., 2013, Kwon et al., 2013)
      yki - CG9581
      anti tag coimmunoprecipitation, peptide massfingerprinting, anti tag western blot
      (Kwon et al., 2013, Kwon et al., 2013)
      yki - CG15117
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Kwon et al., 2013)
      yki - CG31705
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Kwon et al., 2013)
      yki - ci
      anti tag coimmunoprecipitation, anti tag western blot
      (Li et al., 2015)
      yki - Cpsf5
      anti tag coimmunoprecipitation, peptide massfingerprinting, anti tag western blot
      (Kwon et al., 2013, Kwon et al., 2013)
      yki - Cpsf6
      anti tag coimmunoprecipitation, anti tag western blot, peptide massfingerprinting
      (Kwon et al., 2013, Kwon et al., 2013)
      yki - DCAF12
      anti tag coimmunoprecipitation, anti tag western blot
      (Cho et al., 2020)
      yki - dsh
      anti tag coimmunoprecipitation, tag visualisation
      (Varelas et al., 2010)
      yki - EcR
      coimmunoprecipitation, western blot
      (Zhang et al., 2015)
      yki - ed
      anti tag coimmunoprecipitation, anti tag western blot
      (Yue et al., 2012)
      yki - Eip75B
      proximity ligation assay, fluorescence microscopy, anti tag coimmunoprecipitation, western blot
      (Wang et al., 2024)
      yki - EMC1
      anti tag coimmunoprecipitation, peptide massfingerprinting, anti tag western blot
      (Kwon et al., 2013, Kwon et al., 2013)
      yki - ex
      anti tag coimmunoprecipitation, anti tag western blot, pull down, peptide massfingerprinting, Identification by mass spectrometry, anti bait coimmunoprecipitation, western blot
      (Wang et al., 2022, Srivastava et al., 2021, Hu et al., 2016, Kwon et al., 2013, Badouel et al., 2009, Oh et al., 2009)
      yki - foxo
      anti tag coimmunoprecipitation, anti tag western blot, anti bait coimmunoprecipitation, western blot, pull down
      (Sun et al., 2024)
      yki - Fs(2)Ket
      pull down, peptide massfingerprinting
      (Wang et al., 2016)
      yki - Helz
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Kwon et al., 2013)
      yki - Hipk
      enzymatic study, autoradiography, anti tag coimmunoprecipitation, anti tag western blot
      (Steinmetz et al., 2021, Chen and Verheyen, 2012)
      yki - hpo
      anti tag coimmunoprecipitation, peptide massfingerprinting, anti tag western blot
      (Srivastava et al., 2021, Dent et al., 2015, Oh et al., 2009)
      yki - Hsc70-3
      proximity ligation assay, fluorescence microscopy, anti tag coimmunoprecipitation, anti tag western blot, peptide massfingerprinting
      (Yang et al., 2022, Kwon et al., 2013, Kwon et al., 2013)
      yki - hth
      anti tag coimmunoprecipitation, anti tag western blot
      (Srivastava et al., 2021, Peng et al., 2009)
      yki - Inos
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Kwon et al., 2013)
      yki - Ist1
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Kwon et al., 2013)
      yki - Kap-α1
      anti tag coimmunoprecipitation, anti tag western blot, pull down, peptide massfingerprinting
      (Wang et al., 2016)
      yki - Lasp
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Kwon et al., 2013)
      yki - Leash
      anti tag coimmunoprecipitation, anti tag western blot, peptide massfingerprinting
      (Shen et al., 2018, Kwon et al., 2013, Kwon et al., 2013, Kwon et al., 2013)
      yki - Mad
      anti tag coimmunoprecipitation, anti tag western blot, western blot
      (Srivastava et al., 2021, Oh and Irvine, 2011, Alarcón et al., 2009)
      yki - mask
      anti tag coimmunoprecipitation, anti tag western blot
      (Sansores-Garcia et al., 2013, Sidor et al., 2013)
      yki - MED1
      anti bait coimmunoprecipitation, peptide massfingerprinting
      (Oh et al., 2013)
      yki - MED15
      anti bait coimmunoprecipitation, peptide massfingerprinting
      (Oh et al., 2013)
      yki - MED19
      anti bait coimmunoprecipitation, peptide massfingerprinting
      (Oh et al., 2013)
      yki - MED23
      anti bait coimmunoprecipitation, peptide massfingerprinting
      (Oh et al., 2013)
      yki - MED31
      anti bait coimmunoprecipitation, peptide massfingerprinting
      (Oh et al., 2013)
      yki - mesh
      anti tag coimmunoprecipitation, anti tag western blot
      (Chen et al., 2020)
      yki - Mitf
      anti tag coimmunoprecipitation, anti tag western blot
      (Zhang et al., 2024)
      yki - mop
      pull down, peptide massfingerprinting, anti tag coimmunoprecipitation, tandem affinity purification, Identification by mass spectrometry, anti tag western blot
      (Hu et al., 2016, Kwon et al., 2013, Gilbert et al., 2011)
      yki - mor
      anti tag coimmunoprecipitation, anti tag western blot, anti bait coimmunoprecipitation, peptide massfingerprinting, pull down, western blot
      (Srivastava et al., 2021, Oh et al., 2013)
      yki - N
      proximity ligation assay, fluorescence microscopy
      (Wang et al., 2024)
      yki - Ncoa6
      anti tag coimmunoprecipitation, anti tag western blot, pull down, western blot
      (Srivastava et al., 2021, Oh et al., 2014, Qing et al., 2014)
      yki - nerfin-1
      anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based, anti tag western blot
      (Guo et al., 2019, Rhee et al., 2014)
      yki - osa
      anti bait coimmunoprecipitation, peptide massfingerprinting
      (Jin et al., 2013)
      yki - ovo
      anti tag coimmunoprecipitation, anti tag western blot
      (Bohère et al., 2018)
      yki - p53
      anti tag coimmunoprecipitation, anti tag western blot
      (Di Cara et al., 2015)
      yki - Pax
      anti bait coimmunoprecipitation, western blot, anti tag coimmunoprecipitation, anti tag western blot
      (Jiang et al., 2025)
      yki - Pen
      pull down, peptide massfingerprinting
      (Wang et al., 2016)
      yki - Polr2A
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Kwon et al., 2013)
      yki - Prp4k
      enzymatic study, autoradiography
      (Cho et al., 2018)
      yki - Ptp61F
      anti tag coimmunoprecipitation, anti tag western blot, western blot
      (Liu et al., 2022)
      yki - Rae1
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Kwon et al., 2013)
      yki - Rbf
      anti tag coimmunoprecipitation, anti tag western blot, anti bait coimmunoprecipitation
      (Rader et al., 2023)
      yki - RhoGAP15B
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Kwon et al., 2013)
      yki - RpS15Ab
      experimental knowledge based
      (Guruharsha et al., 2011)
      yki - sd
      fluorescent resonance energy transfer, anti tag coimmunoprecipitation, anti tag western blot, surface plasmon resonance, predetermined participant, two hybrid, peptide massfingerprinting, anti bait coimmunoprecipitation, pull down, autoradiography, western blot
      (Sun et al., 2024, Zhai et al., 2024, Mesrouze et al., 2022, Srivastava et al., 2021, Bajpai et al., 2020, Mesrouze et al., 2020, Guo et al., 2019, Cho et al., 2018, Zhang et al., 2017, Li et al., 2015, Ikmi et al., 2014, Cagliero et al., 2013, Guo et al., 2013, Kwon et al., 2013, Oh et al., 2013, Goulev et al., 2008, Wu et al., 2008, Zhang et al., 2008)
      yki - Sec31
      anti tag coimmunoprecipitation, peptide massfingerprinting, anti tag western blot
      (Kwon et al., 2013, Kwon et al., 2013)
      yki - Ssk
      anti tag coimmunoprecipitation, anti tag western blot
      (Chen et al., 2020)
      yki - Strn-Mlck
      anti tag coimmunoprecipitation, anti tag western blot
      (Xu et al., 2018)
      yki - Su(H)
      proximity ligation assay, fluorescence microscopy
      (Wang et al., 2024)
      yki - tai
      coimmunoprecipitation, anti tag western blot, anti tag coimmunoprecipitation, peptide massfingerprinting
      (Zhang et al., 2015, Kwon et al., 2013)
      yki - Tgi
      anti tag coimmunoprecipitation, peptide massfingerprinting, molecular sieving, molecular weight, pull down, anti tag western blot, two hybrid, isothermal titration calorimetry, predetermined participant, Identification by mass spectrometry, nuclear magnetic resonance
      (Srivastava et al., 2021, Vissers et al., 2020, Nyarko, 2018, Guo et al., 2013, Koontz et al., 2013, Kwon et al., 2013)
      yki - Trl
      anti tag coimmunoprecipitation, anti tag western blot, anti bait coimmunoprecipitation, western blot, pull down, peptide massfingerprinting
      (Srivastava et al., 2021, Oh et al., 2013)
      yki - trr
      pull down, western blot
      (Oh et al., 2014)
      yki - tsr
      proximity ligation assay, fluorescence microscopy, anti tag coimmunoprecipitation, western blot, bimolecular fluorescence complementation
      (Skouloudaki et al., 2019)
      yki - Usp7
      anti tag coimmunoprecipitation, anti tag western blot, pull down, western blot
      (Sun et al., 2024, Sun et al., 2019)
      yki - Usp10
      anti tag coimmunoprecipitation, anti tag western blot
      (Gao et al., 2019)
      yki - vg
      anti tag coimmunoprecipitation, anti tag western blot
      (Li et al., 2015)
      yki - Wbp2
      anti tag coimmunoprecipitation, anti tag western blot, peptide massfingerprinting, pull down
      (Srivastava et al., 2021, Hu et al., 2016, Kwon et al., 2013, Zhang et al., 2011)
      yki - wts
      anti tag coimmunoprecipitation, anti tag western blot, enzymatic study, autoradiography, peptide massfingerprinting, western blot, two hybrid, pull down
      (Zhao et al., 2023, Srivastava et al., 2021, Hao et al., 2020, Hu et al., 2016, Hu et al., 2014, Kwon et al., 2013, Sansores-Garcia et al., 2013, Badouel et al., 2009, Oh et al., 2009, Zhang et al., 2009, Wu et al., 2008, Dong et al., 2007, Huang et al., 2005)
      Alleles Reported to Model Human Disease (Disease Ontology) (34 alleles)
      l(2)gl
      Models Based on Experimental Evidence ( 8 )
      Allele
      Disease
      Evidence
      References
      l(2)gl1
      model of  brain cancer
      CEA
      (Betschinger et al., 2006)
      CEA with l(2)gl4
      (Betschinger et al., 2006)
      l(2)gl4
      model of  cancer
      CEA with Ras85DV12.cUa.UAS
      (Fangninou et al., 2023, Liu et al., 2022, Yang et al., 2022)
      CEA with Ras85DV12.UAS
      (Guo et al., 2020, Grifoni et al., 2015, Pagliarini and Xu, 2003)
      CEC with Ras85DV12.UAS
      (Ma et al., 2014)
      CEA
      (Kurelac et al., 2019, Sollazzo et al., 2018, Janic et al., 2010, Woodhouse et al., 2003)
      model of  carcinoma
      CEA
      (Singh et al., 2022, Sollazzo et al., 2022, Rusu et al., 2021, Biehler et al., 2020, Tamori et al., 2010, Bilder et al., 2000)
      CEA with ykiUAS.cOa.Tag:V5
      (Enzo et al., 2015)
      CEA with ykiUAS.cHa
      (Singh et al., 2022)
      CEA with Ras85DV12.UAS
      (Ding et al., 2022, Singh et al., 2022, Ito and Igaki, 2021, Wu et al., 2021, Mishra-Gorur et al., 2019, Zhang et al., 2019, Srivastava et al., 2007, Igaki et al., 2006)
      CEA with Ras85DV12.cUa.UAS
      (Sun et al., 2019)
      model of  brain cancer
      CEA
      (Rossi et al., 2018, Beaucher et al., 2007, Woodhouse et al., 2003)
      CEA with l(2)gl1
      (Betschinger et al., 2006)
      model of  kidney cancer
      CEA
      (Zeng et al., 2010)
      model of  neuroblastoma
      CEA
      (Gateff and Schneiderman, 1974)
      l(2)gl4w3
      model of  carcinoma
      CEA
      (Bilder et al., 2000)
      l(2)glunspecified
      model of  cancer
      CEA with Ras85DV12.UAS
      (Xu et al., 2023)
      l(2)glHMS01522
      model of  carcinoma
      CEA with MycUAS.Tag:HA
      (Sollazzo et al., 2022)
      CEA
      (Singh et al., 2022)
      l(2)glGD4047
      model of  cancer
      CEA with RpL27A1
      (Grifoni et al., 2015)
      CEA
      (Tamori et al., 2016)
      model of  adenocarcinoma
      CEA with Ras85DV12.cUa.UAS
      (Casas-Tintó et al., 2016)
      CEA with wupAf06492, Ras85DV12.cUa.UAS
      (Casas-Tintó et al., 2016)
      model of  carcinoma
      CEA
      (Chatterjee et al., 2023, Jevitt et al., 2021)
      CEA with NCA.UAS.cUa
      (Jevitt et al., 2021)
      l(2)glKK100777
      model of  leukemia
      CEA with Ras85DV12.UAS
      (Arefin et al., 2017)
      model of  brain cancer
      CEA with Ras85DV12.cUa.UAS
      (Manzanero-Ortiz et al., 2021)
      l(2)glHMS01905
      model of  cancer
      CEA
      (Chatterjee et al., 2022)
      Modifiers Based on Experimental Evidence ( 4 )
      yki
      Models Based on Experimental Evidence ( 15 )
      Allele
      Disease
      Evidence
      References
      ykiUAS.cHa
      model of  cancer
      CEA
      (Liu et al., 2022, Yang et al., 2022, Nagarkar et al., 2020, Ziosi et al., 2010)
      CEA with pnutGD1512
      (Gerlach et al., 2018)
      model of  neurodegenerative disease
      CEA
      (Napoletano et al., 2011)
      model of  carcinoma
      CEA
      (Groth et al., 2020, Pérez et al., 2015)
      CEA with l(2)gl4
      (Singh et al., 2022)
      model of  heart disease
      CEA
      (Karekar et al., 2021)
      ykiHMS00041
      model of  carcinoma
      CEA
      (Bhattacharya et al., 2024)
      ykiS168A.UAS.cUa
      model of  cancer
      CEA
      (Jang et al., 2017)
      ykiS168A.UAS.EGFP,Tag:HA
      model of  cancer
      CEA
      (Yeom et al., 2021, Yeom et al., 2020, Bunker et al., 2015)
      ykiUAS.cOa.Tag:V5
      model of  carcinoma
      CEA with l(2)gl4
      (Enzo et al., 2015)
      ykiS168A.UAS.Tag:V5
      model of  cancer
      CEA
      (Hodgson et al., 2021, Santabárbara-Ruiz and Léopold, 2021)
      model of  ovarian cancer
      CEA
      (Hall et al., 2010)
      ykiS111A.S168A.S250A.UAS.Tag:V5
      model of  intestinal cancer
      CEA
      (Xu et al., 2024, Pranoto et al., 2023, Saavedra et al., 2023, Wang et al., 2023, Kim et al., 2022, Bajpai et al., 2020)
      model of  cancer
      CEA
      (Pranoto and Kwon, 2024, Xu et al., 2023, Chen et al., 2022, Ding et al., 2021, Lee et al., 2021)
      CEA with hopTum.UAS
      (Ding et al., 2021)
      model of  neurodegenerative disease
      CEA
      (Napoletano et al., 2011)
      model of  carcinoma
      CEA
      (Singh et al., 2022)
      CEA with scribHMJ21977
      (Snigdha et al., 2021)
      model of  nephrolithiasis
      CEA
      (Xu et al., 2024)
      ykiUAS.cUa
      model of  cancer
      CEA with RhoBTBHMC03199
      (Nguyen et al., 2020)
      CEA with RhoBTBVSH330130
      (Nguyen et al., 2020)
      CEA with Lkb1KK108675
      (Nguyen et al., 2020)
      CEA with Lkb1HMS01351
      (Nguyen et al., 2020)
      CEA with Lkb1GL00019
      (Nguyen et al., 2020)
      CEA with Lkb1VSH330167
      (Nguyen et al., 2020)
      CEA
      (Guo et al., 2020)
      yki3SA.UAS.cUa
      model of  cancer
      CEA
      (Lodge et al., 2021)
      ykiUAS.UTR.Tag:MYC
      model of  cancer
      CEA
      (Guo et al., 2020)
      ykiUAS.UTR-M1.Tag:MYC
      model of  cancer
      CEA
      (Guo et al., 2020)
      ykiUAS.UTR-M2.Tag:MYC
      model of  cancer
      CEA
      (Guo et al., 2020)
      yki3SA.lexAop.Tag:V5
      model of  cancer
      CEA
      (Lee et al., 2021)
      yki3SA.lexAop.Tag:Myr(Unk),GFP
      model of  cancer
      CEA
      (Chen et al., 2022)
      yki3SA.lexAop.sfGFP
      model of  intestinal cancer
      CEA
      (Xu et al., 2024, Saavedra et al., 2023)
      Modifiers Based on Experimental Evidence ( 19 )
      Allele
      Disease
      Interaction
      References
      ykiRNAi.N.UAS
      ameliorates  cancer
      modeled by Ras85DV12.UAS
      (Waghmare et al., 2024)
      exacerbates  Alzheimer's disease 1
      modeled by Hsap\APPAβ42.UAS.cUa
      (Irwin et al., 2020)
      ykiRNAi.C.UAS
      ameliorates  cancer
      modeled by Ras85DV12.UAS
      (Waghmare et al., 2024)
      ykiUAS.cHa
      ameliorates  neurodegenerative disease
      modeled by ft8
      (Napoletano et al., 2011)
      modeled by Zzzz\CAG127Q.UAS.Tag:HA
      (Dubey and Tapadia, 2018)
      modeled by Hsap\HTTQ93.ex1.UAS
      (Dubey and Tapadia, 2018)
      exacerbates  carcinoma
      modeled by hpo3D
      (Pérez et al., 2015)
      model of  carcinoma
      is ameliorated by Atg1UAS.cSa
      (Pérez et al., 2015)
      is exacerbated by Cdk9UAS.ORF
      (Nagarkar et al., 2020)
      is exacerbated by HeximKK104531
      (Nagarkar et al., 2020)
      is exacerbated by NELF-BKK100078
      (Nagarkar et al., 2020)
      is exacerbated by Nelf-AKK101005
      (Nagarkar et al., 2020)
      is exacerbated by bin3KK106875
      (Nagarkar et al., 2020)
      model of  cancer
      is ameliorated by MycGD1419
      (Ziosi et al., 2010)
      is exacerbated by sdUAS.cSa
      (Ziosi et al., 2010)
      is ameliorated by Hsc70-3UAS.ORF.Tag:HA
      (Yang et al., 2022)
      is ameliorated by Xbp1UAS.s.Tag:V5
      (Yang et al., 2022)
      is ameliorated by Ire1UAS.Tag:V5
      (Yang et al., 2022)
      ykiB5
      ameliorates  neurodegenerative disease
      modeled by ft8
      (Napoletano et al., 2011)
      ameliorates  cancer
      modeled by scrib1
      (Bunker et al., 2015)
      modeled by Ras85DV12.UAS, Rox8GD11645
      (Guo et al., 2020)
      ameliorates  brain cancer
      modeled by l(3)mbtGM76
      (Richter et al., 2011)
      ykiNIG.4005R
      ameliorates  cancer
      modeled by shotGL01286
      (Jain et al., 2019)
      ykiHMS00041
      ameliorates  cancer
      modeled by dlg1GD4689, Ras85DV12.UAS
      (Katheder et al., 2017)
      ameliorates  high grade glioma
      modeled by PtenRNAi.Exel.UAS, Ras85DV12.cUa.UAS
      (Gangwani et al., 2020)
      modeled by Pi3K92EUAS.Tag:MYC,Tag:PM(hKRAS), btl::Egfrλ.UAS
      (Vigneswaran et al., 2021)
      DOES NOT ameliorate  cancer
      modeled by HipkUAS.Tag:HA
      (Blaquiere et al., 2018)
      model of  carcinoma
      is ameliorated by AktHMS06047
      (Bhattacharya et al., 2024)
      is ameliorated by ImpL2HMC05809
      (Bhattacharya et al., 2024)
      is ameliorated by Pi3K92EHMC05152
      (Bhattacharya et al., 2024)
      is ameliorated by mTorTED.UAS
      (Bhattacharya et al., 2024)
      ykiGD11187
      ameliorates  intestinal cancer
      modeled by Ptp61FHMS00421
      (Liu et al., 2022)
      ameliorates  kidney disease
      modeled by sdtNIG.15342R
      (Weide et al., 2017)
      modeled by sdtHMS01652
      (Weide et al., 2017)
      ykiKK109756
      ameliorates  colorectal cancer
      modeled by SH3PX1d1
      (Zhang et al., 2019)
      ameliorates  high grade glioma
      modeled by Pi3K92EUAS.Tag:MYC,Tag:PM(hKRAS), btl::Egfrλ.UAS
      (Vigneswaran et al., 2021)
      ykiS168A.UAS.cUa
      model of  cancer
      is ameliorated by lzUAS.cBa
      (Jang et al., 2017)
      is exacerbated by trioUAS.cUa
      (Jang et al., 2017)
      ykiS168A.UAS.EGFP,Tag:HA
      model of  cancer
      is exacerbated by GlnRSKK109443
      (Yeom et al., 2020)
      is ameliorated by AlaRSGD8009
      (Yeom et al., 2020)
      is exacerbated by ArgRSGD14098
      (Yeom et al., 2020)
      is ameliorated by AsnRSGD7064
      (Yeom et al., 2020)
      is ameliorated by AsnRSKK100240
      (Yeom et al., 2020)
      is ameliorated by AspRSGD1339
      (Yeom et al., 2020)
      is exacerbated by CysRSGD13923
      (Yeom et al., 2020)
      is exacerbated by GluProRSGD11635
      (Yeom et al., 2020)
      is ameliorated by GlyRSKK107421
      (Yeom et al., 2020)
      is exacerbated by HisRSKK102374
      (Yeom et al., 2020)
      is exacerbated by LysRSKK101852
      (Yeom et al., 2020)
      is exacerbated by SerRS-mGD12798
      (Yeom et al., 2020)
      is exacerbated by ThrRSGD1342
      (Yeom et al., 2020)
      is exacerbated by TyrRSGD11353
      (Yeom et al., 2020)
      is ameliorated by Ilp8MI00727
      (Yeom et al., 2021)
      is ameliorated by ImpL2GD6004
      (Yeom et al., 2021)
      is ameliorated by Lgr3Δ50
      (Yeom et al., 2021)
      is ameliorated by Ilp8KK112161
      (Yeom et al., 2021)
      exacerbates  cancer
      modeled by HipkUAS.Tag:HA
      (Blaquiere et al., 2018)
      exacerbates  carcinoma
      modeled by NkapGD11807
      (Guo et al., 2024)
      ykiUAS.cOa.Tag:V5
      ameliorates  Alzheimer's disease 1
      modeled by Hsap\APPAβ42.UAS.cUa
      (Irwin et al., 2020)
      ykiS111A.S168A.S250A.UAS.Tag:V5
      model of  intestinal cancer
      is ameliorated by Ilp2UAS.cUa
      (Wang et al., 2023)
      is ameliorated by ImpL2NIG.15009R
      (Wang et al., 2023)
      is ameliorated by mewJF02694
      (Bajpai et al., 2020)
      is ameliorated by rheaHM05161
      (Bajpai et al., 2020)
      is ameliorated by vgTONDU.UAS
      (Bajpai et al., 2020)
      is ameliorated by whdHMS00040
      (Saavedra et al., 2023)
      is exacerbated by zfh2EAB
      (Pranoto et al., 2023)
      is ameliorated by ImpL2HMC05809
      (Pranoto et al., 2023)
      is ameliorated by Mmp1JF01336
      (Pranoto et al., 2023)
      is ameliorated by NDN.UAS.cUa
      (Pranoto et al., 2023)
      is ameliorated by upd2HMS00901
      (Pranoto et al., 2023)
      is ameliorated by upd2HMS00948
      (Pranoto et al., 2023)
      is ameliorated by Pvf1KK112211
      (Xu et al., 2024)
      model of  cancer
      is ameliorated by foxoTM.UAS
      (Lee et al., 2021)
      is ameliorated by panΔN.UAS
      (Lee et al., 2021)
      is ameliorated by wgHMS00794
      (Lee et al., 2021)
      is ameliorated by wgJF01257
      (Lee et al., 2021)
      is ameliorated by AktHMS00007
      (Lee et al., 2021)
      is ameliorated by ITPGD3825
      (Xu et al., 2023)
      is ameliorated by ITPJF01817
      (Xu et al., 2023)
      is ameliorated by ImpL2HMC05809
      (Chen et al., 2022)
      is ameliorated by upd3HMS00646
      (Ding et al., 2021)
      is ameliorated by Stat92EHMS00035
      (Ding et al., 2021)
      is ameliorated by domeGD2612
      (Ding et al., 2021)
      is ameliorated by hopHMS00761
      (Ding et al., 2021)
      ykiJF03119
      ameliorates  Noonan syndrome
      modeled by Hsap\RAF1gof.UAS
      (Yu et al., 2015)
      ykiUAS.cUa
      ameliorates  peroxisome biogenesis disorder 2A
      modeled by Pex5GD14972
      (Pinelli et al., 2024)
      model of  cancer
      is ameliorated by Rox8EY02351
      (Guo et al., 2020)
      ykiRNAi.UAS.cUa
      ameliorates  cancer
      modeled by scribGD11666
      (Ding et al., 2021)
      exacerbates  neurodegenerative disease
      modeled by Zzzz\CAG127Q.UAS.Tag:HA
      (Dubey and Tapadia, 2018)
      modeled by Hsap\HTTQ93.ex1.UAS
      (Dubey and Tapadia, 2018)
      ykiUAS.UTR.Tag:MYC
      model of  cancer
      is ameliorated by Rox8EY02351
      (Guo et al., 2020)
      ykiUAS.UTR-M2.Tag:MYC
      model of  cancer
      is ameliorated by Rox8EY02351
      (Guo et al., 2020)
      yki3SA.lexAop.Tag:Myr(Unk),GFP
      model of  cancer
      is ameliorated by AktΔPH.UAS.Tag:Myr(Unk)
      (Chen et al., 2022)
      is ameliorated by PGRP-SC2UAS.cGa
      (Chen et al., 2022)
      is ameliorated by RelHMS00070
      (Chen et al., 2022)
      yki3SA.lexAop.sfGFP
      model of  intestinal cancer
      is ameliorated by AktΔPH.UAS.Tag:Myr(Unk)
      (Saavedra et al., 2023)
      is ameliorated by REPTORNIG.13624R
      (Saavedra et al., 2023)
      is ameliorated by PvrHMS01662
      (Xu et al., 2024)
      Hsap\YAP1
      Models Based on Experimental Evidence ( 1 )
      Allele
      Disease
      Evidence
      References
      Hsap\YAP1S127A.YAP2.UAS.Tag:FLAG
      model of  ovarian cancer
      CEA
      (Hall et al., 2010)
      Modifiers Based on Experimental Evidence ( 1 )
      Allele
      Disease
      Interaction
      References
      Hsap\YAP1ΔC61.UAS
      ameliorates  Huntington's disease
      modeled by Hsap\HTTGMR.Q120
      (Hoshino et al., 2006)
      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
      Hsap\YAP1UAS.Tag:HA
      P{UAS-hYAP1.HA}
      Hsap\YAP13A.UAS.Tag:HA
      P{UAS-hYAP1.3A.HA}
      Hsap\YAP1ΔC61.UAS
      P{UAS-Hsap\YAP1.DeltaC61}
      Hsap\YAP1ΔC13.UAS
      P{UAS-Hsap\YAP1.DeltaC13}
      Hsap\YAP1ΔC25.UAS
      P{UAS-Hsap\YAP1.DeltaC25}
      Hsap\YAP1UAS.cHa
      P{UAS-YAP1.H}
      Hsap\YAP1S127A.UAS
      P{UAS-YAP.S127A}
      Hsap\YAP1YAP2.UAS.Tag:FLAG
      P{UAS-Flag-YAP2}
      Hsap\YAP1S127A.YAP2.UAS.Tag:FLAG
      P{UAS-Flag-YAP2.S127A}
      Hsap\YAP1S127A.W1W2.UAS.Tag:FLAG
      P{UAS-Flag-YAPS127A/W1W2}
      Hsap\YAP1S94A.S127A.UAS
      P{UAS-YAP-S127A/S94A}
      Hsap\YAP1S94A.UAS
      P{UAS-YAP-S94A}
      Hsap\YAP1W1W2.UAS
      P{UAS-YAP.W1W2}
      Hsap\YAP1UAS.cZa
      P{UAS-YAP1.Z}
      Hsap\YAP1UAS.cIa
      P{UAS-YAP1.I}
      Hsap\YAP1S127A.UAS.cIa
      P{UAS-YAP1.S127A}
      Hsap\LLGL1UAS.cGa
      P{UAS-Hugl1.G}
      Selected Drosophila transgenes
      Allele
      Transgene
      Publicly Available Stocks
      yki3SA.UAS.cUa
      P{UAS-yki.3SA.U}
      ykiUAS.UTR.Tag:MYC
      P{UAS-Myc-yki.UTR}
      ykiUAS.UTR-M1.Tag:MYC
      P{UAS-Myc-yki.UTR-M1}
      ykiUAS.UTR-M2.Tag:MYC
      P{UAS-Myc-yki.UTR-M2}
      yki3SA.lexAop.Tag:V5
      P{lexAop-yki.3SA.V5}
      yki3SA.lexAop.Tag:Myr(Unk),GFP
      P{lexAop-yki.3SA.MyrGFP}
      yki3SA.lexAop.sfGFP
      P{lexAop-yki.S3A.sfGFP}
      ykiUAS.cHa
      P{UAS-yki}
      w*; P{UAS-yki}3/TM3, Sb1
      ykiS168A.UAS.EGFP,Tag:HA
      P{UAS-yki.S168A.GFP.HA}
      y1 w*; P{UAS-yki.S168A.GFP.HA}10-7-Y
      ykiUAS.cOa.Tag:V5
      P{UAS-yki.V5.O}
      w*; P{UAS-yki.V5.O}attP2
      ykiS168A.UAS.Tag:V5
      P{UAS-yki.S168A.V5}
      w*; P{UAS-yki.S168A.V5}attP2
      ykiS111A.S168A.S250A.UAS.Tag:V5
      P{UAS-yki.S111A.S168A.S250A.V5}
      w*; P{UAS-yki.S111A.S168A.S250A.V5}attP2
      ykiS168A.UAS.cUa
      P{UAS-yki.S168A.U}
      ykiUAS.cUa
      P{UAS-yki.U}
      RNAi constructs available
      Allele
      Transgene
      Publicly Available Stocks
      ykiRNAi.13xlexAop.cZa
      P{13xlexAop-yki.RNAi.Z}
      ykiGD11187
      P{GD11187}
      w1118; P{GD11187}v40497/TM3
      ykiKK109756
      P{KK109756}
      P{KK109756}VIE-260B
      ykiRNAi.N.UAS
      P{UAS-yki.RNAi.N}
      ykiRNAi.C.UAS
      P{UAS-yki.RNAi.C}
      ykiJF03119
      P{TRiP.JF03119}
      y1 v1; P{TRiP.JF03119}attP2/TM3, Sb1
      ykiNIG.4005R
      P{NIG.4005R}
      ykiHMS00041
      P{TRiP.HMS00041}
      y1 v1; P{TRiP.HMS00041}attP2
      ykiRNAi.UAS.cUa
      P{UAS-yki.RNAi.U}
      ykiRNAi.UASp.cLa
      P{UASp-yki.RNAi.L}
      l(2)glRNAi.UAS.cBa
      P{UAS-l(2)gl.RNAi.B}
      l(2)glRNAi.UAS.cUa
      P{UAS-l(2)gl.RNAi.U}
      l(2)glGD4047
      P{GD4047}
      w1118; P{GD4047}v51247
      l(2)glJF01073
      P{TRiP.JF01073}
      y1 v1; P{TRiP.JF01073}attP2
      l(2)glJF01553
      P{TRiP.JF01553}
      y1 v1; P{TRiP.JF01553}attP2
      l(2)glKK100777
      P{KK100777}
      P{KK100777}VIE-260B
      l(2)glHMS01522
      P{TRiP.HMS01522}
      y1 sc* v1 sev21; P{TRiP.HMS01522}attP2
      l(2)glGL00645
      P{TRiP.GL00645}
      y1 sc* v1 sev21; P{TRiP.GL00645}attP40
      l(2)glHMS01905
      P{TRiP.HMS01905}
      y1 v1; P{TRiP.HMS01905}attP40
      Selected Drosophila classical alleles
      Allele
      Allele class
      Mutagen
      Publicly Available Stocks
      ykiAcidA
      loss of function allele
      CRISPR/Cas9
      w*; P{neoFRT}42D ykiAcidA/CyO, P{Dfd-EYFP.wBmC}2
      ykiB5
      loss of function allele
      l(2)gl4 ykiB5/CyO
      l(2)gl275
      loss of function allele
      spontaneous
      l(2)glunspecified
      l(2)gl4
      amorphic allele - genetic evidence
      spontaneous
      y1 w67c23; l(2)gl4/CyO, y+
      l(2)gl4w3
      w*; l(2)gl4w3 P{neoFRT}40A P{FRT(whs)}G13/CyO
      l(2)gl1
      spontaneous
      l(2)glts3
      ethyl methanesulfonate
      y1 w*; l(2)glts3 cn1 speck1/CyO, y+
      l(2)glGX7w-
      amorphic allele - molecular evidence
      ends-out gene targeting
      References (10)