migration in the Drosophila gastrula. Curr Biol
14:659–667. https://doi.org/10.1016/j.cub.
2004.03.058
8. Crawford JM, Harden N, Leung T et al (1998)
Cellularization in Drosophila melanogaster is
disrupted by the inhibition of rho activity and
the activation of Cdc42 function. Dev Biol
204:151–164.
https://doi.org/10.1006/
dbio.1998.9061
9. Barrett K, Leptin M, Settleman J (1997) The
Rho GTPase and a putative RhoGEF mediate a
signaling pathway for the cell shape changes in
Drosophila gastrulation. Cell 91:905–915
10. Magie CR, Meyer MR, Gorsuch MS, Parkhurst
SM (1999) Mutations in the Rho1 small
GTPase disrupt morphogenesis and segmentation during early Drosophila development.
Development 126:5353–5364
11. Ni J-Q, Zhou R, Czech B et al (2011) A
genome-scale shRNA resource for transgenic
RNAi
in
Drosophila.
Nat
Methods
8:405–407.
https://doi.org/10.1038/
nmeth.1592
12. Fristrom D, Fristrom JW (1993) The metamorphic development of the adult epidermis.
In: Bate M, Alfonso Martinez Arias A (eds) The
development of Drosophila melanogaster. Cold
Spring Harbor Laboratory Press, Plainview, pp
843–897
13. Pastor-Pareja JC, Grawe F, Martı ´n-Blanco E,
Garcı ´a-Bellido A (2004) Invasive cell behavior
during Drosophila imaginal disc eversion is
mediated by the JNK signaling cascade. Dev
Cell 7:387–399. https://doi.org/10.1016/j.
devcel.2004.07.022
14. Aldaz S, Escudero LM, Freeman M (2010)
Live imaging of Drosophila imaginal disc
development. Proc Natl Acad Sci U S A
107:14217–14222.
https://doi.org/10.
1073/pnas.1008623107
15. Aldaz S, Escudero LM, Freeman M (2013)
Dual role of myosin II during Drosophila
imaginal disc metamorphosis. Nat Commun
4:1761–1710.
https://doi.org/10.1038/
ncomms2763
16. Proag A, Monier B, Suzanne M (2019) Physical and functional cell-matrix uncoupling in a
developing tissue under tension. Development
146:dev172577–9.
https://doi.org/10.
1242/dev.172577
17. Murray MJ (2015) Drosophila models of
metastasis. AIMS Genet 2(1):25–53. https://
doi.org/10.3934/genet.2015.1.25
18. Ishimaru S, Ueda R, Hinohara Y et al (2004)
PVR plays a critical role via JNK activation in
thorax closure during Drosophila metamorphosis. EMBO J 23:3984–3994. https://doi.
org/10.1038/sj.emboj.7600417
19. Srivastava A, Pastor-Pareja JC, Igaki T et al
(2007) Basement membrane remodeling is
essential for Drosophila disc eversion and
tumor invasion. Proc Natl Acad Sci U S A
104:2721–2726. https://doi.org/10.1073/
pnas.0611666104
20. Martı ´n-Blanco E, Pastor-Pareja JC, GarciaBellido A (2000) JNK and decapentaplegic signaling control adhesiveness and cytoskeleton
dynamics during thorax closure in Drosophila.
Proc Natl Acad Sci U S A 97:7888–7893
21. Manhire-Heath R, Golenkina S, Saint R, Murray MJ (2013) Netrin-dependent downregulation of Frazzled/DCC is required for the
dissociation of the peripodial epithelium in
Drosophila. Nat Commun 4:1–10. https://
doi.org/10.1038/ncomms3790
22. Tripura C, Chandrika N-P, Susmitha V-N et al
(2011) Regulation and activity of JNK signaling in the wing disc peripodial membrane during adult morphogenesis in Drosophila. Int J
Dev Biol 55:583–590. https://doi.org/10.
1387/ijdb.103275ct
23. Dietzl G, Chen D, Schnorrer F et al (2007) A
genome-wide transgenic RNAi library for conditional gene inactivation in Drosophila.
Nature 448:151–156. https://doi.org/10.
1038/nature05954
24. Gibson MC, Lehman DA, Schubiger G (2002)
Lumenal transmission of decapentaplegic in
Drosophila imaginal discs. Dev Cell 3:451–460
25. Pallavi SK, Shashidhara LS (2003) Egfr/Ras
pathway mediates interactions between peripodial and disc proper cells in Drosophila wing
discs. Development 130:4931–4941. https://
doi.org/10.1242/dev.00719
26. Larsen C, Franch-Marro X, Hartenstein V et al
(2006) An efficient promoter trap for detection
of patterned gene expression and subsequent
functional analysis in Drosophila. Proc Natl
Acad Sci U S A 103:17813–17817. https://
doi.org/10.1073/pnas.0607652103
27. Calleja M, Moreno E, Pelaz S, Morata G
(1996) Visualization of gene expression in living adult Drosophila. Science 274:252–255
28. McGuire SE, Mao Z, Davis RL (2004) Spatiotemporal gene expression targeting with the
TARGET and gene-switch systems in Drosophila. Sci STKE 2004:pl6. https://doi.org/
10.1126/stke.2202004pl6
29. Green EW, Fedele G, Giorgini F, Kyriacou CP
(2014) A Drosophila RNAi collection is subject to dominant phenotypic effects. Nat Methods 11:222–223. https://doi.org/10.1038/
nmeth.2856
30. Vissers JHA, Manning SA, Kulkarni A, Harvey
KF (2016) A Drosophila RNAi library modulates Hippo pathway-dependent tissue growth.
Using Drosophila Wing Eversion to Study EMT
133
14:659–667. https://doi.org/10.1016/j.cub.
2004.03.058
8. Crawford JM, Harden N, Leung T et al (1998)
Cellularization in Drosophila melanogaster is
disrupted by the inhibition of rho activity and
the activation of Cdc42 function. Dev Biol
204:151–164.
https://doi.org/10.1006/
dbio.1998.9061
9. Barrett K, Leptin M, Settleman J (1997) The
Rho GTPase and a putative RhoGEF mediate a
signaling pathway for the cell shape changes in
Drosophila gastrulation. Cell 91:905–915
10. Magie CR, Meyer MR, Gorsuch MS, Parkhurst
SM (1999) Mutations in the Rho1 small
GTPase disrupt morphogenesis and segmentation during early Drosophila development.
Development 126:5353–5364
11. Ni J-Q, Zhou R, Czech B et al (2011) A
genome-scale shRNA resource for transgenic
RNAi
in
Drosophila.
Nat
Methods
8:405–407.
https://doi.org/10.1038/
nmeth.1592
12. Fristrom D, Fristrom JW (1993) The metamorphic development of the adult epidermis.
In: Bate M, Alfonso Martinez Arias A (eds) The
development of Drosophila melanogaster. Cold
Spring Harbor Laboratory Press, Plainview, pp
843–897
13. Pastor-Pareja JC, Grawe F, Martı ´n-Blanco E,
Garcı ´a-Bellido A (2004) Invasive cell behavior
during Drosophila imaginal disc eversion is
mediated by the JNK signaling cascade. Dev
Cell 7:387–399. https://doi.org/10.1016/j.
devcel.2004.07.022
14. Aldaz S, Escudero LM, Freeman M (2010)
Live imaging of Drosophila imaginal disc
development. Proc Natl Acad Sci U S A
107:14217–14222.
https://doi.org/10.
1073/pnas.1008623107
15. Aldaz S, Escudero LM, Freeman M (2013)
Dual role of myosin II during Drosophila
imaginal disc metamorphosis. Nat Commun
4:1761–1710.
https://doi.org/10.1038/
ncomms2763
16. Proag A, Monier B, Suzanne M (2019) Physical and functional cell-matrix uncoupling in a
developing tissue under tension. Development
146:dev172577–9.
https://doi.org/10.
1242/dev.172577
17. Murray MJ (2015) Drosophila models of
metastasis. AIMS Genet 2(1):25–53. https://
doi.org/10.3934/genet.2015.1.25
18. Ishimaru S, Ueda R, Hinohara Y et al (2004)
PVR plays a critical role via JNK activation in
thorax closure during Drosophila metamorphosis. EMBO J 23:3984–3994. https://doi.
org/10.1038/sj.emboj.7600417
19. Srivastava A, Pastor-Pareja JC, Igaki T et al
(2007) Basement membrane remodeling is
essential for Drosophila disc eversion and
tumor invasion. Proc Natl Acad Sci U S A
104:2721–2726. https://doi.org/10.1073/
pnas.0611666104
20. Martı ´n-Blanco E, Pastor-Pareja JC, GarciaBellido A (2000) JNK and decapentaplegic signaling control adhesiveness and cytoskeleton
dynamics during thorax closure in Drosophila.
Proc Natl Acad Sci U S A 97:7888–7893
21. Manhire-Heath R, Golenkina S, Saint R, Murray MJ (2013) Netrin-dependent downregulation of Frazzled/DCC is required for the
dissociation of the peripodial epithelium in
Drosophila. Nat Commun 4:1–10. https://
doi.org/10.1038/ncomms3790
22. Tripura C, Chandrika N-P, Susmitha V-N et al
(2011) Regulation and activity of JNK signaling in the wing disc peripodial membrane during adult morphogenesis in Drosophila. Int J
Dev Biol 55:583–590. https://doi.org/10.
1387/ijdb.103275ct
23. Dietzl G, Chen D, Schnorrer F et al (2007) A
genome-wide transgenic RNAi library for conditional gene inactivation in Drosophila.
Nature 448:151–156. https://doi.org/10.
1038/nature05954
24. Gibson MC, Lehman DA, Schubiger G (2002)
Lumenal transmission of decapentaplegic in
Drosophila imaginal discs. Dev Cell 3:451–460
25. Pallavi SK, Shashidhara LS (2003) Egfr/Ras
pathway mediates interactions between peripodial and disc proper cells in Drosophila wing
discs. Development 130:4931–4941. https://
doi.org/10.1242/dev.00719
26. Larsen C, Franch-Marro X, Hartenstein V et al
(2006) An efficient promoter trap for detection
of patterned gene expression and subsequent
functional analysis in Drosophila. Proc Natl
Acad Sci U S A 103:17813–17817. https://
doi.org/10.1073/pnas.0607652103
27. Calleja M, Moreno E, Pelaz S, Morata G
(1996) Visualization of gene expression in living adult Drosophila. Science 274:252–255
28. McGuire SE, Mao Z, Davis RL (2004) Spatiotemporal gene expression targeting with the
TARGET and gene-switch systems in Drosophila. Sci STKE 2004:pl6. https://doi.org/
10.1126/stke.2202004pl6
29. Green EW, Fedele G, Giorgini F, Kyriacou CP
(2014) A Drosophila RNAi collection is subject to dominant phenotypic effects. Nat Methods 11:222–223. https://doi.org/10.1038/
nmeth.2856
30. Vissers JHA, Manning SA, Kulkarni A, Harvey
KF (2016) A Drosophila RNAi library modulates Hippo pathway-dependent tissue growth.
Using Drosophila Wing Eversion to Study EMT
133