publication list – University of Copenhagen

Stephen Cohen

Publication list

H-index 70; >15K citations


Foronda, D., Weng, R., Verma, P., Chen. Y.-W. & Cohen S.M. (submitted) Coordination of Insulin and Notch pathway activities by microRNA miR-305 mediates adaptive homeostasis in the intestinal stem cells of the Drosophila gut


Hong, X., Nguyen T.H., Chen Q., Zhang, R., Voorhoeve, P.M. and Cohen, S.M. (2014). Opposing activities of the Ras and Hippo pathways converge on regulation of YAP protein turnover. EMBO J in press

Nguyen T.H., Hong, X., Tan, S., Chen Q., Chan,L., Fivaz.M., Cohen, S.M. and Voorhoeve, P.M. (2014) Viral small T oncoproteins transform cells by alleviating Hippo pathway-mediated inhibition of the YAP proto-oncogene. Cell Reports 8, 707-713. doi: 10.1016/j.celrep.2014.06.062

Herranz, H., Weng, R., and Cohen, S.M. (2014) Cross-talk between epithelial and mesenchymal tissues in tumorigenesis and imaginal disc development. Curr Biol 24, 1476–1484. doi: 10.1016/j.cub.2014.05.043

Garg, D., and Cohen, S.M.  (2014) miRNAs and aging: a genetic perspective. Ageing Research Reviews; doi: 10.1016/j.arr.2014.04.001.

Cohen, S.M. (2014) Everything old is new again: lincRNAs make proteins. EMBO J 33, 937-938; doi: 10.1002/embj.201488303.


Kugler, J-M., Chen, Y-W., Weng, R.,  and Cohen, S.M. (2013) miRNAs act maternally to confer robustness to Drosophila primordial germ cell formation. G3, doi: 10.1534/g3.113.007591

Kugler, J-M., Verma, P., Chen, Y-W., Weng, R.,  & Cohen, SM (2013) miR-989 is required for border cell migration in the Drosophila ovary. PloS One. doi: 10.1371/journal.pone.0067075

Zhang, W. and Cohen, S.M.  (2013) The Hippo pathway acts via p53 and microRNAs to control proliferation and proapoptotic gene expression during tissue growth. Biology Open 2, 822-828; doi:10.1242/bio.20134317  

Weng, R., Chin, J.S.R., Yew, J.Y., Bushati, N. & Cohen, SM (2013) miR-124 controls male reproductive success in Drosophila. eLife 2013;2:e00640

Zhang, W., Hietakangas, V., Wee, S., Lim, R.  Gunaratne, J., and Cohen, S.M.  (2013) ER stress potentiates insulin resistance through PERK-mediated FOXO phosphorylation. Genes Dev 27: 441-449.


Vodala, S., Pescatore, S., Rodriguez, J., Buescher, M., Chen, Y., Weng, R., Cohen, S.M. and Rosbash, M. (2012) The Oscillating miRNA 959-964 cluster impacts Drosophila feeding time and other circadian outputs. Cell Metabolism 16, 601-612.

Herranz, H., Hong, X., Nguyen T.H., Voorhoeve, P.M. and Cohen, S.M.  (2012) Oncogenic cooperation between SOCS family proteins and EGFR identified using a Drosophila epithelial transformation model. Genes Dev 26, 1602-1611.

Aw, S.  and Cohen, S.M.  (2012) Time is of the essence: microRNAs and age-associated neurodegeneration. Cell Research, doi:10.1038/cr.2012

Herranz, H., Hong, X. and Cohen, S.M.  (2012) Mutual repression by bantam miRNA and Capicua links the EGFR/MAPK and Hippo pathways in growth control. Current Biol. 22, 651-657. doi:10.1016/j.cub.2012.02.050

Weng, R. and Cohen, S.M. (2012) Drosophila miR-124 regulates neuroblast proliferation through its target anachronism. Development, 139, 1427-1434.

Ge, W., Chen Y.-W. Weng, R., Lim, S.-F. Buescher,M., Zhang, R. and S.M. Cohen (2012) Overlapping functions of microRNAs in control of apoptosis during Drosophila embryogenesis. Cell Death and Differentiation 19, 839-846. doi:10.1038/cdd.2011.161


Zhang, W., Thompson, B.J. Hietakangas, V., and SM Cohen (2011) MAPK/ERK signaling regulates insulin sensitivity to control glucose metabolism in Drosophila. PLoS Genet, 7: e1002429. doi:10.1371/journal.pgen.1002429

Szuplewski, S., Kugler. J-M., Lim, S.F, Verma, P., Chen, Y-W. and S.M. Cohen (2011) MicroRNA transgene overexpression complements deficiency-based modifier screens in Drosophila. Genetics, 190:617-626  doi:10.1534/genetics.111.136689

Becam, I., Rafel, N., Hong, X., Cohen, S.M. and M Milan (2011) Notch mediated repression of bantam miRNA contributes to boundary formation in the Drosophila wing. Development 138:3781-9.

Chen, Y.-W. Weng, R. and Cohen, S.M. (2011) Protocols for use of homologous recombination gene-targeting to produce microRNA mutants in Drosophila Methods in Molecular Biology, 732: 99-120.


Varghese, J., Lim, S., and SM Cohen (2010) Drosophila miR-14 regulates insulin production and metabolism through its target sugarbabe. Genes Dev. 24: 2748-2753.

Milán M. and S.M. Cohen (2010) Lateral inhibition: intermittent long range Notch signaling to achieve robustness. Current Biol 20:R802-R804.

Cohen SM (2010) Editorial.  Seminars in Cell and Developmental Biology 21, 727.

Bryk, B., Hahn, K., Cohen, S.M. and Teleman A.A. (2010) MAP4K3 regulates body size and metabolism in Drosophila. Dev Biol, 344,150-7.

Herranz, H. and SM Cohen  (2010) microRNAs and gene regulatory networks: managing the impact of noise in biological systems. Genes Dev 24, 1339-1344.

Hilgers,V., Bushati, N., and Cohen, SM (2010) Drosophila microRNAs 263a/b confer robustness during development by protecting nascent sense organs from apoptosis. PLoS Biology 15;8(6):e1000396

Herranz, H., Hong, X., Pérez, L., Ferreira, A., Olivieri, D., Cohen, S.M. and Milán, M (2010) A role of Dcr-1 in regulating Myc protein levels in the Drosophila wing. EMBO J  29,  1688-98.

Bennett EP, Chen YW, Schwientek T, Mandel U, Schjoldager KB, Cohen SM, Clausen H. (2010) Rescue of Drosophila Melanogaster l(2)35Aa lethality is only mediated by polypeptide GalNAc-transferase pgant35A, but not by the evolutionary conserved human ortholog GalNAc-transferase-T11. Glycoconj J. 27:435-44. Epub 2010 Apr 27.

Cohen, S.M. (2010) microRNAs in CNS development and neurodegeneration: insights from Drosophila genetics. In Macro-roles for microRNAs in the life and death of neurons. IPSEN Foundation Springer (B de Strooper and Y Christensen eds) pp 69-77.


Cohen, S.M. (2009) Use of microRNA sponges to explore tissue specific microRNA functions in vivo. Nature Methods. 6, 873-874.

Hong, X., Hammell, M. Ambros, V and Cohen, S.M. (2009) Identification of microRNA targets by immunopurification of Ago1 RNPs: selection for a distinct class of targets. PNAS 106, 15085-90

Hietakangas, V., and Cohen, S.M. (2009) Regulation of tissue growth through nutrient sensing. Annual Reviews of  Genetics 43, 389–410.

Szuplewski, S., Sandmann, T., Hietakangas, V. and Cohen, S.M. (2009) Drosophila Minus is required for cell proliferation and influences Cyclin E turnover. Genes Dev 17, 1998-2003

Weng, R., Chen, Y.W. Bushati, N. Cliffe A. and Cohen, S.M. (2009) Recombinase-Mediated Cassette Exchange provides a versatile platform for gene targeting: knockout of miR-31b. Genetics 183, 399-402

Pizette, S., Rabouille, C., Cohen, S.M. and Thérond, P.  (2009) Glycosphingolipids control the extracellular gradient of the Drosophila EGFR ligand Gurken. Development 136, 551-561.


Hietakangas, V., and Cohen, S.M. (2008) TOR complex 2 is needed for cell cycle progression and anchorage-independent growth of MCF7 and PC3 tumor cells. BMC Cancer 8, 282

Neumüller, R.A., Betschinger, J., Fischer, A., Bushati, N., Poernbacher, I., Mechtler, K., Cohen, S.M., and J.A. Knoblich (2008) Mei-P26 regulates microRNAs and cell growth in the Drosophila ovarian stem cell lineage. Nature 454, 241-245.

Bushati, N., Stark, A., Brennecke, J., and Cohen, S.M. (2008) Temporal reciprocity of microRNAs and their targets during the maternal to zygotic transition in Drosophila. Current Biology 18, 501-506.

Stark, A., Bushati, N., Jan, C., Hodges, E., Brennecke, J., Bartel, D., Cohen, S.M. and Kellis, M. (2008) Drosophila hox iab-4 produces functional sense and anti-sense miRNAs from a single hairpin. Genes Dev 22, 8-13.

Teleman, A.A., Hietakangas, V., Sayadian, A.C. and Cohen, S.M. (2008) Nutritional control of protein biosynthetic capacity: convergent regulation of Myc by the FOXO and TORC1 branches of the Insulin/TOR pathway. Cell Metabolism 7, 21-32.

Bushati, N & Cohen, S.M. (2008) microRNAs in neurodegeneration. Curr Opin Neurobiol 18, 292-296

Hietakangas, V. & Cohen, S.M. (2008) TORCing up metabolic control in the brain. Cell Metabolism 7, 357-358.


Sandmann, T. and Cohen, S.M. (2007) Identification of novel Drosophila melanogaster microRNAs. PLoS One 2(11): e1265. doi:10.1371/journal.pone.0001265

Karres, J.S., Hilgers, V. Carrera, I. Treisman, J. and Cohen, S.M. (2007) The conserved microRNA miR-8 tunes Atrophin levels to prevent neurodegeneration in Drosophila. Cell 131, 136–145.

Varghese, J. & Cohen, S.M. (2007) microRNA miR-14 acts in a feed-forward loop controlling steroid hormone signaling in Drosophila. Genes Dev 21, 2277-2282.

Easow, G., Teleman,A.A.,  and Cohen, S.M. (2007) Isolation of microRNA targets by miRNP immunopurification. RNA 13, 1198-1204.

Chen, Y.-W., Pedersen, J.W., Wandall, H.H., Levery, S.B., Pizette, S. Clausen, H., & Cohen, S.M. (2007) Glycosphingolipids with extended sugar chain have specialized functions in development and behavior of Drosophila Dev Biol 306, 736-49.

Hietakangas, V. & Cohen, S.M. (2007) Re-evaluating AKT regulation: role of TOR complex 2 in tissue growth. Genes Dev. 21, 632-637.

Hufnagel, L., Teleman,A.A., Rouault , H., Cohen, S.M. & Shraiman B.I.,  (2007) On the mechanism of wing size determination in fly development. PNAS 104, 3835-3840.

Bushati, N & Cohen, S.M. (2007) microRNA functions. Annual Reviews of Cell and Developmental Biology  23:175-205.


Cohen S.M., Brennecke, J. & Stark, A. (2006) Peak selection: a new role for microRNAs in neurogenesis. Genes Dev 20: 2769-277

Cohen S.M. & Brennecke, J. (2006) Mixed messages in early development. Science 312:65-6.

Hufnagel, L., Kreuger, J., Cohen, S.M. & Shraiman B.I. (2006) On the role of glypicans in the process of morphogen gradient formation. Dev Biol 300, 512-522.

Jinek, M., Chen Y.-W., Clausen H., Cohen, S.M.  & Conti, E. (2006) Structural insights into the Notch-modifying glycosyltransferase Fringe. Nature Str Mol Biol 13, 945-946.

Thompson, B.J. & Cohen, S.M. (2006) The Hippo pathway regulates the bantam microRNA to control cell proliferation and apoptosis in Drosophila. Cell 126, 767-774.

Teleman, A.A., Maitra S. & Cohen S.M. (2006) Drosophila lacking microRNA miR-278 are defective in energy homeostasis Genes Dev. 20,417-422.

Rehwinkel, J., Natalin, P., Stark, A., Brennecke, J., Cohen, S.M. & Izaurralde, E . (2006) Genome-wide analysis of mRNAs regulated by Drosha and Argonaute proteins in Drosophila Mol Cell Biol 26, 2965-75.


Stark, A., Brennecke, J., Bushati, N. Russell, R.B. & Cohen, S.M. (2005) Animal microRNAs confer robustness to gene expression and have a significant impact on 3’UTR evolution. Cell 123,1133-1146.

Thompson, B.J., Mathieu,J., Sung,H.-H., Loeser,E. Rørth,P. & Cohen, S.M. (2005) Tumour suppressor properties of the ESCRT-II complex component, Vps25, in Drosophila Dev Cell 9, 711-720.

Brennecke, J., Stark, A., & Cohen S.M. (2005) Not miR-ly muscular: microRNAs and muscle development. Genes Dev 19:2261-4.

Weihe, U., Milán, M. & Cohen, S.M. (2005) Drosophila limb development. in: Comprehensive Molecular Insect Science. Volume 1.  L. Gilbert, K. Iatrou and S. Gill (editors) pp 305-341,

Staudt, N., Molitor, A., Somogyi, K., Mata, J., Curado, S., Eulenberg, K., Meise, M., Siegmund, T., Häder, T., Hilfiker, A., Brönner, G., Ephrussi, A., Rørth, P., Cohen, S.M., Fellert, S., Chung, H-R., Piepenburg, O., Schäfer, U., Jäckle, H. & Vorbrüggen, G. (2005 ) Gain-of-Function Screen for Genes That Affect Drosophila Muscle Pattern Formation. PloS Genet, 1(4): e55

Teleman,A.A., Chen, Y.-W. & Cohen S.M. (2005) 4E-BP functions as a metabolic brake used under stress conditions but not during normal growth. Genes Dev. 19: 1844-1848.

Teleman,A.A., Chen, Y.-W. & Cohen S.M. (2005) Drosophila Melted modulates FOXO and TOR activity. Dev Cell 9, 271–281.

Brennecke, J., Stark, A., Russell, R.B. & Cohen, S.M. (2005) Principles of microRNA/target recognition. PLoS Biol. 2005 Mar;3(3):E85. epub 2005 Feb15

Neubueser, D., Warren, J.T., Gilbert, L. I., & Cohen, S.M. (2005) molting defective is required for ecdysone biosynthesis. Dev Biol. 280, 362-372.

Collins R.T. & Cohen S.M. (2005) A genetic screen in Drosophila to identity novel components of the Hedgehog signaling pathway. Genetics published March 2, 2005 DOI: 10.1534/genetics.104.039420

Perez, L., Milan M., Bray, S. & Cohen, S.M. (2005) Ligand-binding and signaling properties of the Ax[M1] form of Notch. Mech Dev. 122, 479-486.

Mikeladze-Dvali, T., Wernet, M.F., Pistillo, D., Mazzoni, E.O., Teleman, A.A., Chen, Y.-W., Cohen, SM & Desplan, C. (2005) The Growth Regulators warts/lats and melted Interact in a Bistable Loop to Specify Opposite Fates in Drosophila R8 Photoreceptors. Cell 122, 775-787.

Wandall, H., Pizette, S., Pedersen, J.W., Eichert, H., Levery, S.B., Mandel, U., Clausen, H. & Cohen, S.M. (2005) Egghead and brainiac are essential for glycosphingolipid biosynthesis in vivo. J Biol Chem 280:4858-63. Epub Dec 15 2004.

Milan M, Perez L, Cohen SM (2005) Boundary formation in the Drosophila wing: Functional dissection of Capricious and Tartan. Dev Dynamics  233, 804-810.


Weihe, U., Dorfman, R., Wernet, M. F., S. M. Cohen & Milán, M. (2004) Proximal-distal subdivision of Drosophila legs and wings: the elbow-no ocelli gene complex. Development 131: 767-74.

Emerald, B.S. & Cohen S.M. (2004) The homeotic selector gene Antennapedia establishes leg identity by repressing antennal identity in Drosophila. Dev. Biol. 267: 462-72.

Hipfner, D.R., Keller N. & Cohen, S.M. (2004) Slik Sterile-20 kinase regulates Moesin activity to promote epithelial integrity during tissue growth. Genes Dev 18: 2243-2248.

Milan M., Pham T.T. & Cohen S.M. (2004) Osa modulates the expression of Apterous target genes in the Drosophila wing. Mech Dev. 121:491-7.

Kreuger, J., Perez, L., Giraldez, A. J. & Cohen, S.M. (2004) Opposing activities of Dally-like glypican at high and low levels of Wingless morphogen activity. Dev Cell 7:503-512.

Hipfner, D. & Cohen, S.M. (2004) Connecting proliferation and apoptosis in development and disease. Nature Reviews Cell Mol Biol 5, 805-815.


Brennecke, J., Hipfner, D.R. Stark, A., Russell, R & Cohen, S.M. (2003) bantam encodes a developmentally regulated microRNA that controls cell proliferation and regulates the pro-apoptotic gene hid in Drosophila. Cell 113: 25-36.

Milán, M. & Cohen, S.M. (2003) A re-evaluation of the contributions of Apterous and Notch to the Dorsal-Ventral lineage restriction boundary in the Drosophila wing. Development 130: 553-562.

Emerald, B.S., Curtiss, J., Mlodzik, M. & Cohen S.M. (2003) Distal antenna and distal antenna related encode nuclear proteins containing pipsqueak motifs involved in antenna development in Drosophila Development 130: 1171-1180.

Wandall, H.H., Pedersen, J.W., Park, C., Levery, S.B., Pizette, S., Cohen, S.M., Schwientek, T. & Clausen, H. (2003) Drosophila Egghead Encodes a b1,4Mannosyltransferase Predicted to Form the Immediate Precursor Glycosphingolipid Substrate for Brainiac. J Biol. Chem. 278:1411-1414.

Hipfner, D.R. & Cohen, S.M. (2003) The Drosophila Sterile-20 Kinase, slik controls cell proliferation and apoptosis during imaginal disc development. PLoS Biol. 2003 Nov;1(2):E35. Epub 2003 Nov 17.

Stark, A., Brennecke, J., Russell, R & Cohen, S.M. (2003) Identification of microRNA targets. PLoS Biol. 2003 Dec;1(3):E60. Epub 2003 Oct 13.

Giraldez, A.J. & Cohen S.M. (2003) Wingless and Notch signaling provide cell survival cues and control cell proliferation during wing development. Development 130: 6533-6543.

Collins, R.T. & Cohen S.M. (2003) The secret life of Smoothened. Dev Cell 5: 823-824. (preview)

Cohen, S.M. (2003) Long-range signalling by touch. Nature 426: 503-4. (news & views)

Brennecke, J. & Cohen, S.M. (2003) Towards a complete description of the microRNA complement of animal genomes. Genome Biology 4:228.


Britton, J.S., Lockwood, W.K., Li, L., Cohen, S.M. & Edgar, B.A. (2002) Drosophila's Insulin/PI3-Kinase Pathway Coordinates Cellular Metabolism with Nutritional Conditions. Developmental Cell 2, 239-249.

Wu, J. & Cohen, S.M. (2002)  Repression of Teashirt marks the initiation of wing development. Development 129, 2411-2418.

Giraldez, A.J. Copley, R.R. & Cohen, S.M. (2002) Regulation of Wingless gradient formation by Notum: a novel secreted enzyme that modifies proteoglycans. Developmental Cell 2, 667-676.

Milán, M., Pérez, L. & Cohen, S.M. (2002) Short-range cell interactions and cell survival in the Drosophila wing. Developmental Cell 2, 797-805.

Giraldez, A.J., Pérez, L.  & Cohen, S.M. (2002) A naturally occurring alternative product of the mastermind locus that represses Notch signaling. Mech Dev. 115, 101-105.

Hipfner, D.R., Weigmann, K., & Cohen, S.M. (2002) The Bantam gene product regulates Drosophila growth. Genetics 161: 1527-1537.

Schwientek, T., Keck, B., Levery, S.B., Jensen, M.A., Pedersen, J.W., Wandall, H., Stroud, M., Cohen, S.M., Amado, M. & H. Clausen (2002) The Drosophila Gene Brainiac Encodes a Glycosyltransferase Putatively Involved in Glycosphingolipid Synthesis. J Biol. Chem. 277:32421-9.


Kerber, B., Monge, I., Mueller, M., Mitchell, P.J. & Cohen, S.M. (2001) The AP-2 transcription factor is required for joint formation and cell survival in Drosophila leg development. Development 128, 1231-1238

Milán, M. Weihe, U., Tiong, S., Bender, W. & Cohen, S.M. (2001) msh specifies dorsal cell fate in the Drosophila wing. Development 128, 3263-3268.

Milán, M. Weihe, U., Pérez, L. & Cohen, S.M. (2001) The LRR-proteins Capricious and Tartan mediate cell interactions during boundary formation in the Drosophila wing. Cell, 106, 785-794.

Weihe, U., Milán, M. & Cohen, S.M. (2001) Regulation of Apterous activity in Drosophila wing development. Development 128: 4615-4622.

Teleman A.A., Strigini M, & Cohen, S.M. (2001) Shaping morphogen gradients. Cell 105, 559-562.

Starling Emerald, B.  & Cohen, S.M. (2001) Limb development: getting down to the ground state. Current Biology 11, R1025–R1027.


Strigini, M. & Cohen, S.M. (2000) Wingless gradient formation in the Drosophila wing imaginal disc. Current Biology 10, 293-300.

Wu, J. & Cohen, S.M. (2000) Proximal distal axis formation in the Drosophila leg: Distinct functions of Teashirt and Homothorax in the proximal leg. Mech Devel 94, 47-56.

Milán, M. & Cohen, S.M. (2000) Temporal regulation of Apterous activity during development of the Drosophila wing. Development 127, 3069-3078.

Milán, M. & Cohen, S.M. (2000) Subdividing cell populations in the developing limbs of Drosophila to progressively smaller functional units: wing veins and leg segments. Dev Biol 217, 1-9.

Brückner, K., Perez, L., Clausen, H. & Cohen, S.M. (2000) Glycosyltransferase activity of Fringe modulates Notch-Delta interactions. Nature 406, 411-415.

Denef, N., Neubüser, D. Perez, L. & Cohen, S.M. (2000) Hedgehog induces opposite changes in turnover and subcellular localization of Patched and Smoothened. Cell 102, 521-531.

Teleman A.A. & Cohen, S.M. (2000) Dpp gradient formation in the Drosophila wing imaginal disc. Cell 103, 971-980


Wu, J. & Cohen, S.M. (1999) Proximal distal axis formation in the Drosophila leg: Primary subdivision into proximal and distal domains by Homothorax, Teashirt and Distal-less expression.  Development 126, 109-117.

Milán, M. & Cohen, S.M. (1999) Regulation of LIM-homeodomain protein activity in vivo: A tetrameric complex of dLDB and Apterous confers Apterous activity and the capacity for regulation by the antagonist dLMO. Molecular Cell 4, 267-273.

Weigmann, K. & Cohen, S.M. (1999) Lineage tracing cells born in different domains along the PD axis of the developing Drosophila leg. Development 126, 3823-3830.

Milán, M. & Cohen, S.M. (1999) Notch signaling is not sufficient to define the affinity boundary between dorsal and ventral compartments. Molecular Cell 4, 1073-1078.

Cohen, S.M. (1999) Decapentaplegic morphogen gradient in Drosophila wing imaginal disc. in Cell Surface Proteoglycans in Signalling and Development (A. Lander, Nakato, H., Selleck, S.B., Turnbull, J.E., Coath, C. eds) pp 188-193

Strigini M, & Cohen, S.M. (1999) Formation of morphogen gradients in the Drosophila wing. Sem Cell Dev. Biol 10, 335-344.

Hipfner, D.R. & Cohen, S.M. (1999) New growth factors for imaginal discs. Bioessays 21, 718-720.


Rorth, P., Szabo, K., Bailey, A. Laverty, T., Rehm, J., Rubin, G. M., Weigmann, K., Milan, M., Benes, V., Ansorge, W., &  Cohen, S.M. (1998) Systematic Gain-of-Function Genetics in Drosophila. Development 125, 1049-1057.

Neumann, C.J. & Cohen, S.M. (1998) Boundary formation in the Drosophila wing: the POU-domain protein Nubbin attenuates Notch activity. Science 281, 409-413.

Milán, M., Diaz-Benjumea, F.J. & Cohen, S.M. (1998) An LMO protein regulates LIM-homeodomain activity in Drosophila wing development: a possible molecular model for LMO oncogenes. Genes Dev. 12, 2912-2920.

Lecuit, T. & Cohen, S.M. (1998) Dpp receptor levels contribute to shaping the Dpp morphogen gradient in the Drosophila wing imaginal disc. Development 125, 4901-4907.

Royet, J., Bouwmeester, T. & Cohen, S.M. (1998) Notchless encodes a novel WD40-repeat-containing protein that modulates Notch signaling activity. EMBO J. 17: 7351-7360


Averof, M. & Cohen, S.M. (1997) Evolutionary origin of insects wings from ancestral gills. Nature 385, 627-630.

Neumann, C.J. & Cohen, S.M. (1997) Long-range action of Wingless organizes the D/V axis of the Drosophila wing. Development 124, 871-880

Neumann, C.J. & Cohen, S.M. (1997) Morphogens and pattern formation. Bioessays  19, 721-729.

Voie A.-M. & Cohen S.M. (1997) Transgenic Drosophila. in Methods in Cell Biology (J. Celis ed.)

Wimmer, E.A., Cohen S.M., Jäckle, H. & Desplan C. (1997) buttonhead does not contribute to a combinatorial code proposed for Drosophila head development. Development 124, 1509-1517.

Lecuit, T. & Cohen, S.M. (1997) Proximal-Distal axis formation in the Drosophila leg. Nature 388, 139-145.

Weigmann, K., Cohen, S.M. & Lehner, C. (1997) Cell cycle progression, growth and patterning in imaginal discs despite inhibition of cell division after inactivation of Drosophila Cdc2 kinase. Development 124, 3555-3563.

Strigini, M. & Cohen, S.M. (1997) A Hedgehog activity gradient contributes to AP axial patterning of the Drosophila wing. Development 124, 4697-4705.


Neumann, C.J.  &  Cohen, S.M. (1996) Sternopleural is a regulatory mutation of wingless with both dominant and recessive effects on larval development of Drosophila melanogaster.  Genetics 142, 1147-1155.

Neumann, C.J. & Cohen, S.M. (1996) Distinct mitogenic and cell fate specification functions of wingless in different regions of the wing. Development 122, 1782-1789.

Ng, M., Diaz-Benjumea, F.J., Vincent, J.-P., Wu, J. & Cohen, S.M. (1996) Specification of the wing by localized expression of Wingless protein. Nature 381, 316-318.

Lecuit, T., Brook, W.J., Ng, M., Calleja, M., Sun, H., & Cohen, S.M. (1996) Two distinct mechanisms for long-range patterning by Decapentaplegic in the Drosophila wing. Nature 381, 387-393.

Wiersdorff, V., Lecuit, T., Cohen, S.M, & Mlodzik, M. (1996) Mad acts downstream of Dpp receptors, and reveals a differential requirement for dpp signaling in initiation and propagation of morphogenesis in the Drosophila eye. Development 122, 2153-2162.

Brook, W.J. & Cohen, S.M. (1996) Antagonistic interactions between Wingless and Decapentaplegic responsible for dorsal-ventral pattern in the Drosophila leg. Science 273, 1373-1377.

Neumann, C.J. & Cohen, S.M. (1996) A hierarchy of cross-regulation involving Notch, wingless, vestigial and cut organizes the dorsal/ventral axis of the Drosophila wing. Development 122, 3477-3485.

Eaton, S. & Cohen, S.M. (1996) More than one way to skin a b-cat. Trends Cell Biol. 6, 287-290.

Cohen, S.M. (1996) Controlling growth of the wing: vestigial integrates signals from the compartment boundaries. Bioessays 18, 855-858.

Brook, W.J, Diaz-Benjumea, F.J. & Cohen, S.M. (1996) Organizing spatial pattern in limb development. Annual Reviews of Cell and Developmental Biology 12: 161-180


Ng, M., Diaz-Benjumea, F.J. & Cohen, S.M. (1995) nubbin encodes a POU-domain protein required for proximal-distal patterning of the Drosophila  wing. Development 121, 589-599.

Lepage, T., Cohen, S.M., Diaz-Benjumea, F.J. & Parkhurst, S. (1995) Signal transduction by cAMP-dependent protein kinase A in Drosophila limb patterning. Nature 373, 711-715.

Simmonds, A.J., Brook, W. J., Cohen, S. M. & Bell, J. B. (1995) Distinct functions for engrailed and invected in anterior-posterior patterning in the Drosophila wing. Nature 376, 424-427.

Wimmer, E.A., Simpson-Brose, M, Cohen, S.M., Desplan, C. & Jäckle, H. (1995) Trans- and cis-acting requirements for expression of the head gap gene buttonhead. Mech Devel. 53: 235-245.

Diaz-Benjumea, F.J. & Cohen, S.M. (1995) Serrate signals through Notch to establish a Wingless-dependent organizer at the dorsal/ventral compartment boundary of the Drosophila wing. Development, 121, 4215-4225


Diaz-Benjumea, F.J. & Cohen, S.M. (1994) wingless acts through the shaggy/ zeste-white 3 kinase to direct dorsal-ventral axis formation in the Drosophila leg. Development 120, 1661-1670.

Diaz-Benjumea, F.J., Cohen B. & Cohen, S.M. (1994) Cell interaction between compartments establishes the proximal-distal axis of Drosophila legs. Nature 372, 175-179.

Cohen, S.M.  & Hyman, A.A. (1994) When is a determinant a determinant? Current Biology 4, 420-422.


Cohen, B., Simcox, A.A. & Cohen S.M. (1993) Allocation of thoracic imaginal discs in the Drosophila embryo. Development 117, 597-608.

O'Hara, E., Cohen B., Cohen, S.M. & McGinnis, W. (1993) Distal-less is a downstream gene of Deformed required for ventral maxillary identity. Development 117, 847-856.

Thüringer, F., Cohen, S.M. & Bienz, M. (1993) Dissection of an indirect autoregulatory response of a homeotic Drosophila gene. EMBO J. 12, 2419-2430.

Diaz-Benjumea, F.J. & Cohen, S.M. (1993) Interaction between dorsal and ventral cells in the imaginal disc directs wing development in Drosophila. Cell 75, 741-752.

Wimmer, E.A., Jäckle, H., Pfeifle, C. & Cohen, S.M. (1993) A Drosophila homolog of human Sp1 is a head-specific segmentation gene. Nature 366, 690-694.

Cohen, S.M. (1993) Imaginal disc development. in Development of Drosophila  Vol. II (M. Bate & A. Martinez Arias eds.) Cold Spring Harbor Press. Cold Spring Harbor N.Y. pp 747-841.

Cohen, S.M. & DiNardo, S. (1993) wingless: from embryo to adult. Trends Genet. 9, 189-192.


Cohen, B., McGuffin, M.E., Pfeifle, C., Segal, D. & Cohen, S.M. (1992) apterous: a gene required for imaginal disc development encodes a member of the LIM family of developmental regulatory proteins. Genes Dev. 6, 715-729

Vachon, G., Cohen, B., Pfeifle, C., McGuffin, M.E., Botas, J. & Cohen, S.M. (1992) Homeotic genes of the Bithorax complex repress limb development in the abdomen of the Drosophila embryo through the target gene Distal-less.  Cell 71, 437-450.

Cohen, B & Cohen, S.M. (1992) Double labelling of mRNA and proteins in Drosophila embryos. in Non-radioactive labelling and detection of Biomolecules (R. Kessler, ed.) Springer Verlag Heidelberg pp. 382-392.


Cohen, B., Wimmer, E. & Cohen, S.M. (1991) Early development of the leg and wing primordia in the Drosophila embryo. Mech. Devel. 33, 229-240.

Simcox, A.A., Hersperger, E., Shearn, A., Whittle, J.R.S. & Cohen, S.M. (1991) Establishment of imaginal discs and histoblast nests in Drosophila. Mech Devel. 34, 11-20.

Cohen, S.M. & Jürgens, G. (1991) Drosophila headlines. Trends Genet. 7, 267-272.


Cohen, S.M. (1990) Specification of limb development in the Drosophila embryo by positional cues from segmentation genes. Nature 343, 173-177.

Cohen, S.M. & Jürgens, G (1990) Mediation of Drosophila head development by gap-like segmentation genes. Nature 346, 482-485.


Cohen, S.M., Brönner, G., Küttner, F., Jürgens, G. & Jäckle, H (1989) Distal-less encodes a homeodomain protein required for limb development in Drosophila. Nature 338, 432-434.

Cohen, S.M. & Jürgens, G. (1989a) Proximal-distal pattern formation in Drosophila:  cell autonomous requirement for Distal-less gene activity in limb development. EMBO J. 8, 2045-2055.

Cohen, S.M. & Jürgens, G. (1989b) Proximal-distal pattern formation in Drosophila:  graded requirement for Distal-less gene activity during limb development. Roux´s Archiv. Dev. Biol. 198, 157-169.


Knecht, D.A, Cohen, S.M., Loomis, W.F. & Lodish, H.F.(1986) Developmental regulation of Dictyostelium actin gene fusions carried on low-copy and high-copy transformation vectors. Mol. Cell. Biol. 6, 3973-3983.

Cohen, S.M., Knecht, D.A, Lodish, H.F. & Loomis, W.F. (1986) DNA sequences required for expression of a Dictyostelium actin gene. EMBO J. 5, 3361-3366.

Cohen, S.M., Nauber, U., Schuh, R., Seifert, E. & Jäckle, H. (1988) Phenocopy production by anti-sense RNA identifies genes required for pattern formation in the Drosophila embryo. in Anti-sense RNA  (D. Melton, ed.) Cold Spring Harbor Press. Cold Spring Harbor N.Y. pp 65-69.

Cappello, J., Cohen, S.M., Handelsman, K & Lodish, H.F. (1986) Structure and regulated transcription of DIRS-1: a novel Dictyostelium discoideum  transposable element. in Genetics Development and Evolution (J.P. Gustafson, G.L.Stebbins and F.J.Ayala) eds. Plenum Publ. Corp. N.Y. pp 235-251.


Gorbsky, G., Cohen, S.M., Shida, H., Giudice, G.J. & Steinberg, M.S.(1985)  Isolation of the non-glycosylated proteins of desmosomes and immunolocalization of a third plaque protein: Desmoplakin III.  Proc. Natl. Acad. Sci. USA. 82, 810-814.

Cappello, J., Handelsman, K., Cohen, S.M. & Lodish, H.F.(1985) Structure and regulated transcription of DIRS-1: an apparent retrotransposon of Dictyostelium discoideum. Cold Spring Harbor Symp. Quant. Biol. 50, 759-767.


Giudice, G.J., Cohen, S.M., Patel, N. & Steinberg, M.S. (1984)  Immunological comparison of desmosomal components from several bovine tissues.  J. Cell. Biochem. 26, 35-45.

Cappello, J., Cohen, S.M. & Lodish, H.F. (1984) Dictyostelium transposable element DIRS-1 preferentially inserts into DIRS-1 sequences. Mol. Cell. Biol. 4, 2207-2213.

Cohen, S.M., Cappello, J. & Lodish, H.F. (1984) Transcription of the Dictyostelium  transposable element DIRS-1. Mol. Cell. Biol. 4, 2332-2340.

Cohen, S.M., Cappello, J., Zuker, C & Lodish, H.F. (1984) Transcription of DIRS-1: an unusual Dictyostelium transposable element. in Molecular Biology of Development  (R.Firtel and E. Davidson) eds. Alan R. Lis Inc. N.Y. pp.491-508.


Cohen, S.M., Gorbsky, G. & Steinberg, M.S. (1983) Immunochemical characterization of related families of glycoproteins in desmosomes. J. Biol. Chem. 258, 2621-2627.

Gorbsky, G., Cohen, S.M., & Steinberg, M.S. (1983) Desmosomal antigens are not recognized by the majority of Pemphigus autoimmune sera. J. Invest. Dermatol. 80, 475-480.