1.
Perez-Vale KZ, Peifer M. Orchestrating morphogenesis: building the body plan by cell shape changes
and movements. Development. 2020 Sep 11; 147(17). https://doi.org/10.1242/dev.191049 PMID:
32917667
2.
Fuchs Y, Steller H. Programmed cell death in animal development and disease. Cell. 2011 Nov 11; 147
(4):742–58. https://doi.org/10.1016/j.cell.2011.10.033 PMID: 22078876
3.
Day SJ, Lawrence PA. Measuring dimensions: the regulation of size and shape. Development. 2000
Jul; 127(14):2977–87. https://doi.org/10.1242/dev.127.14.2977 PMID: 10862736
4.
Tada M, Heisenberg CP. Convergent extension: using collective cell migration and cell intercalation to
shape embryos. Development. 2012 Nov; 139(21):3897–904. https://doi.org/10.1242/dev.073007
PMID: 23048180
5.
Xie T, Spradling AC. A niche maintaining germ line stem cells in the Drosophila ovary. Science. 2000
Oct 13; 290(5490):328–30. https://doi.org/10.1126/science.290.5490.328 PMID: 11030649
6.
Xie T, Spradling AC. decapentaplegic is essential for the maintenance and division of germline stem
cells in the Drosophila ovary. Cell. 1998 Jul 24; 94(2):251–60. https://doi.org/10.1016/s0092-8674(00)
81424-5 PMID: 9695953
PLOS Genetics | https://doi.org/10.1371/journal.pgen.1010684 March 27, 2023
18 / 21
PLOS GENETICS
Shaping GSC-niche by Sas-Ptp10D via JNK-mediated apoptosis
7.
Spradling A, Drummond-Barbosa D, Kai T. Stem cells find their niche. Nature. 2001 Nov 1; 414
(6859):98–104. https://doi.org/10.1038/35102160 PMID: 11689954
8.
Song X, Zhu CH, Doan C, Xie T. Germline stem cells anchored by adherens junctions in the Drosophila
ovary niches. Science. 2002 Jun 7; 296(5574):1855–7. https://doi.org/10.1126/science.1069871 PMID:
12052957
9.
Irizarry J, Stathopoulos A. FGF signaling supports Drosophila fertility by regulating development of
ovarian muscle tissues. Dev Biol. 2015 Aug 1; 404(1):1–13. https://doi.org/10.1016/j.ydbio.2015.04.023
PMID: 25958090
10.
Gilboa L. Organizing stem cell units in the Drosophila ovary. Curr Opin Genet Dev. 2015 Jun; 32:31–6.
https://doi.org/10.1016/j.gde.2015.01.005 PMID: 25703842
11.
Sahut-Barnola I, Godt D, Laski FA, Couderc JL. Drosophila ovary morphogenesis: analysis of terminal
filament formation and identification of a gene required for this process. Dev Biol. 1995 Jul; 170(1):127–
35. https://doi.org/10.1006/dbio.1995.1201 PMID: 7601303
12.
Cohen ED, Mariol MC, Wallace RM, Weyers J, Kamberov YG, Pradel J, et al. DWnt4 regulates cell
movement and focal adhesion kinase during Drosophila ovarian morphogenesis. Dev Cell. 2002 Apr; 2
(4):437–48. https://doi.org/10.1016/s1534-5807(02)00142-9 PMID: 11970894
13.
Song X, Call GB, Kirilly D, Xie T. Notch signaling controls germline stem cell niche formation in the Drosophila ovary. Development. 2007 Mar; 134(6):1071–80. https://doi.org/10.1242/dev.003392 PMID:
17287246
14.
Morata G. Cell competition: A historical perspective. Dev Biol. 2021 Aug; 476:33–40. https://doi.org/10.
1016/j.ydbio.2021.02.012 PMID: 33775694
15.
Baker NE. Emerging mechanisms of cell competition. Nat Rev Genet. 2020 Nov; 21(11):683–97.
https://doi.org/10.1038/s41576-020-0262-8 PMID: 32778819
16.
Johnston LA. Competitive interactions between cells: death, growth, and geography. Science. 2009 Jun
26; 324(5935):1679–82. https://doi.org/10.1126/science.1163862 PMID: 19556501
17.
Nagata R, Igaki T. Cell competition: Emerging mechanisms to eliminate neighbors. Dev Growth Differ.
2018 Dec; 60(9):522–30. https://doi.org/10.1111/dgd.12575 PMID: 30443922
18.
Kanda H, Igaki T. Mechanism of tumor-suppressive cell competition in flies. Cancer Sci. 2020 Oct; 111
(10):3409–15. https://doi.org/10.1111/cas.14575 PMID: 32677169
19.
Vishwakarma M, Piddini E. Outcompeting cancer. Nat Rev Cancer. 2020 Mar; 20(3):187–98. https://
doi.org/10.1038/s41568-019-0231-8 PMID: 31932757
20.
Yamamoto M, Ohsawa S, Kunimasa K, Igaki T. The ligand Sas and its receptor PTP10D drive tumoursuppressive cell competition. Nature. 2017 Feb 9; 542(7640):246–50. https://doi.org/10.1038/
nature21033 PMID: 28092921
21.
Jeon M, Nguyen H, Bahri S, Zinn K. Redundancy and compensation in axon guidance: genetic analysis
of the Drosophila Ptp10D/Ptp4E receptor tyrosine phosphatase subfamily. Neural Dev. 2008 Jan 31;
3:3. https://doi.org/10.1186/1749-8104-3-3 PMID: 18237413
22.
Igaki T, Pastor-Pareja JC, Aonuma H, Miura M, Xu T. Intrinsic tumor suppression and epithelial maintenance by endocytic activation of Eiger/TNF signaling in Drosophila. Dev Cell. 2009 Mar; 16(3):458–65.
https://doi.org/10.1016/j.devcel.2009.01.002 PMID: 19289090
23.
Brumby AM, Richardson HE. scribble mutants cooperate with oncogenic Ras or Notch to cause neoplastic overgrowth in Drosophila. EMBO J. 2003 Nov 3; 22(21):5769–79. https://doi.org/10.1093/emboj/
cdg548 PMID: 14592975
24.
Schonbaum CP, Organ EL, Qu S, Cavener DR. The Drosophila melanogaster stranded at second (sas)
gene encodes a putative epidermal cell surface receptor required for larval development. Dev Biol.
1992 Jun; 151(2):431–45. https://doi.org/10.1016/0012-1606(92)90183-h PMID: 1339334
25.
Sun Q, Schindelholz B, Knirr M, Schmid A, Zinn K. Complex genetic interactions among four receptor
tyrosine phosphatases regulate axon guidance in Drosophila. Mol Cell Neurosci. 2001 Feb; 17(2):274–
91. https://doi.org/10.1006/mcne.2000.0939 PMID: 11178866
26.
Sun Q, Bahri S, Schmid A, Chia W, Zinn K. Receptor tyrosine phosphatases regulate axon guidance
across the midline of the Drosophila embryo. Development. 2000 Feb; 127(4):801–12. https://doi.org/
10.1242/dev.127.4.801 PMID: 10648238
27.
Kai T, Spradling A. An empty Drosophila stem cell niche reactivates the proliferation of ectopic cells.
Proc Natl Acad Sci U S A. 2003 Apr 15; 100(8):4633–8. https://doi.org/10.1073/pnas.0830856100
PMID: 12676994
28.
Jeon M, Zinn K. Receptor tyrosine phosphatases control tracheal tube geometries through negative
regulation of Egfr signaling. Development. 2009 Sep; 136(18):3121–9. https://doi.org/10.1242/dev.
033597 PMID: 19675131
PLOS Genetics | https://doi.org/10.1371/journal.pgen.1010684 March 27, 2023
19 / 21
PLOS GENETICS
Shaping GSC-niche by Sas-Ptp10D via JNK-mediated apoptosis
29.
Zhu CH, Xie T. Clonal expansion of ovarian germline stem cells during niche formation in Drosophila.
Development. 2003 Jun; 130(12):2579–88. https://doi.org/10.1242/dev.00499 PMID: 12736203
30.
Song X, Wong MD, Kawase E, Xi R, Ding BC, McCarthy JJ, et al. Bmp signals from niche cells directly
repress transcription of a differentiation-promoting gene, bag of marbles, in germline stem cells in the
Drosophila ovary. Development. 2004 Mar; 131(6):1353–64. https://doi.org/10.1242/dev.01026 PMID:
14973291
31.
Cabrera GR, Godt D, Fang PY, Couderc JL, Laski FA. Expression pattern of Gal4 enhancer trap insertions into the bric a brac locus generated by P element replacement. Genesis. 2002 Sep-Oct; 34(1–
2):62–5.
32.
Hsu TH, Yang CY, Yeh TH, Huang YC, Wang TW, Yu JY. The Hippo pathway acts downstream of the
Hedgehog signaling to regulate follicle stem cell maintenance in the Drosophila ovary. Sci Rep. 2017
Jun 30; 7(1):4480. https://doi.org/10.1038/s41598-017-04052-6 PMID: 28667262
33.
McGuire SE, Le PT, Osborn AJ, Matsumoto K, Davis RL. Spatiotemporal rescue of memory dysfunction
in Drosophila. Science. 2003 Dec 5; 302(5651):1765–8. https://doi.org/10.1126/science.1089035
PMID: 14657498
34.
Sarov M, Barz C, Jambor H, Hein MY, Schmied C, Suchold D, et al. A genome-wide resource for the
analysis of protein localisation in Drosophila. Elife. 2016 Feb 20; 5:e12068. https://doi.org/10.7554/
eLife.12068 PMID: 26896675
35.
Moreno E, Valon L, Levillayer F, Levayer R. Competition for Space Induces Cell Elimination through
Compaction-Driven ERK Downregulation. Curr Biol. 2019 Jan 7; 29(1):23–34 e8. https://doi.org/10.
1016/j.cub.2018.11.007 PMID: 30554899
36.
Siegrist SE, Haque NS, Chen CH, Hay BA, Hariharan IK. Inactivation of both Foxo and reaper promotes
long-term adult neurogenesis in Drosophila. Curr Biol. 2010 Apr 13; 20(7):643–8. https://doi.org/10.
1016/j.cub.2010.01.060 PMID: 20346676
37.
Xue D, Horvitz HR. Inhibition of the Caenorhabditis elegans cell-death protease CED-3 by a CED-3
cleavage site in baculovirus p35 protein. Nature. 1995 Sep 21; 377(6546):248–51. https://doi.org/10.
1038/377248a0 PMID: 7675111
38.
Meier P, Silke J, Leevers SJ, Evan GI. The Drosophila caspase DRONC is regulated by DIAP1. EMBO
J. 2000 Feb 15; 19(4):598–611. https://doi.org/10.1093/emboj/19.4.598 PMID: 10675329
39.
Shen YH, Godlewski J, Zhu J, Sathyanarayana P, Leaner V, Birrer MJ, et al. Cross-talk between JNK/
SAPK and ERK/MAPK pathways: sustained activation of JNK blocks ERK activation by mitogenic factors. J Biol Chem. 2003 Jul 18; 278(29):26715–21. https://doi.org/10.1074/jbc.M303264200 PMID:
12738796
40.
Katagiri C, Masuda K, Urano T, Yamashita K, Araki Y, Kikuchi K, et al. Phosphorylation of Ser-446
determines stability of MKP-7. J Biol Chem. 2005 Apr 15; 280(15):14716–22. https://doi.org/10.1074/
jbc.M500200200 PMID: 15689616
41.
Dorstyn L, Colussi PA, Quinn LM, Richardson H, Kumar S. DRONC, an ecdysone-inducible Drosophila
caspase. Proc Natl Acad Sci U S A. 1999 Apr 13; 96(8):4307–12. https://doi.org/10.1073/pnas.96.8.
4307 PMID: 10200258
42.
Kondo S, Senoo-Matsuda N, Hiromi Y, Miura M. DRONC coordinates cell death and compensatory proliferation. Mol Cell Biol. 2006 Oct; 26(19):7258–68. https://doi.org/10.1128/MCB.00183-06 PMID:
16980627
43.
Moreno E, Yan M, Basler K. Evolution of TNF signaling mechanisms: JNK-dependent apoptosis triggered by Eiger, the Drosophila homolog of the TNF superfamily. Curr Biol. 2002 Jul 23; 12(14):1263–8.
https://doi.org/10.1016/s0960-9822(02)00954-5 PMID: 12176339
44.
Kuranaga E, Kanuka H, Igaki T, Sawamoto K, Ichijo H, Okano H, et al. Reaper-mediated inhibition of
DIAP1-induced DTRAF1 degradation results in activation of JNK in Drosophila. Nat Cell Biol. 2002 Sep;
4(9):705–10. https://doi.org/10.1038/ncb842 PMID: 12198495
45.
Shlevkov E, Morata G. A dp53/JNK-dependant feedback amplification loop is essential for the apoptotic
response to stress in Drosophila. Cell Death Differ. 2012 Mar; 19(3):451–60. https://doi.org/10.1038/
cdd.2011.113 PMID: 21886179
46.
Kurada P, White K. Ras promotes cell survival in Drosophila by downregulating hid expression. Cell.
1998 Oct 30; 95(3):319–29. https://doi.org/10.1016/s0092-8674(00)81764-x PMID: 9814703
47.
Bergmann A, Agapite J, McCall K, Steller H. The Drosophila gene hid is a direct molecular target of
Ras-dependent survival signaling. Cell. 1998 Oct 30; 95(3):331–41. https://doi.org/10.1016/s00928674(00)81765-1 PMID: 9814704
48.
Decotto E, Spradling AC. The Drosophila ovarian and testis stem cell niches: similar somatic stem cells
and signals. Dev Cell. 2005 Oct; 9(4):501–10. https://doi.org/10.1016/j.devcel.2005.08.012 PMID:
16198292
PLOS Genetics | https://doi.org/10.1371/journal.pgen.1010684 March 27, 2023
20 / 21
PLOS GENETICS
Shaping GSC-niche by Sas-Ptp10D via JNK-mediated apoptosis
49.
Kirilly D, Wang S, Xie T. Self-maintained escort cells form a germline stem cell differentiation niche.
Development. 2011 Dec; 138(23):5087–97. https://doi.org/10.1242/dev.067850 PMID: 22031542
50.
Hay BA, Wolff T, Rubin GM. Expression of baculovirus P35 prevents cell death in Drosophila. Development. 1994 Aug; 120(8):2121–9. https://doi.org/10.1242/dev.120.8.2121 PMID: 7925015
51.
Martin-Blanco E, Gampel A, Ring J, Virdee K, Kirov N, Tolkovsky AM, et al. puckered encodes a phosphatase that mediates a feedback loop regulating JNK activity during dorsal closure in Drosophila.
Genes Dev. 1998 Feb 15; 12(4):557–70. https://doi.org/10.1101/gad.12.4.557 PMID: 9472024
52.
Igaki T, Kanda H, Yamamoto-Goto Y, Kanuka H, Kuranaga E, Aigaki T, et al. Eiger, a TNF superfamily
ligand that triggers the Drosophila JNK pathway. EMBO J. 2002 Jun 17; 21(12):3009–18. https://doi.
org/10.1093/emboj/cdf306 PMID: 12065414
53.
Freeman M. Reiterative use of the EGF receptor triggers differentiation of all cell types in the Drosophila
eye. Cell. 1996 Nov 15; 87(4):651–60. https://doi.org/10.1016/s0092-8674(00)81385-9 PMID: 8929534
54.
Barolo S, Carver LA, Posakony JW. GFP and beta-galactosidase transformation vectors for promoter/
enhancer analysis in Drosophila. BioTechniques. 2000 Oct; 29(4):726, 8, 30, 32. https://doi.org/10.
2144/00294bm10 PMID: 11056799
55.
Ring JM, Martinez Arias A. puckered, a gene involved in position-specific cell differentiation in the dorsal
epidermis of the Drosophila larva. Dev Suppl. 1993:251–9. PMID: 8049480
56.
Schindelin J, Arganda-Carreras I, Frise E, Kaynig V, Longair M, Pietzsch T, et al. Fiji: an open-source
platform for biological-image analysis. Nat Methods. 2012 Jun 28; 9(7):676–82. https://doi.org/10.1038/
nmeth.2019 PMID: 22743772
57.
Kanda Y. Investigation of the freely available easy-to-use software ’EZR’ for medical statistics. Bone
Marrow Transplant. 2013 Mar; 48(3):452–8. https://doi.org/10.1038/bmt.2012.244 PMID: 23208313
PLOS Genetics | https://doi.org/10.1371/journal.pgen.1010684 March 27, 2023
21 / 21
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