49. Adult Drosophila Lack Hematopoiesis but Rely on a Blood Cell Reservoir at the Respiratory Epithelia to Relay Infection Signals to Surrounding Tissues. Sanchez Bosch P, Makhijani K, Herboso L, Gold KS, Baginsky R, Woodcock KJ, Alexander B, Kukar K, Corcoran S, Jacobs T, Ouyang D, Wong C, Ramond EJV, Rhiner C, Moreno E, Lemaitre B, Geissmann F, Brückner K. Dev Cell. 2019 Dec 16;51(6):787-803.e5. doi: 10.1016/j.devcel.2019.10.017. Epub 2019 Nov 14. PMID: 31735669

Times cited: 1

Altmetric score: 17


48. HIF-transcribed p53 chaperones HIF-1α.Madan E, Parker TM, Pelham CJ, Palma AM, Peixoto ML, Nagane M, Chandaria A, Tomás AR, Canas-Marques R, Henriques V, Galzerano A, Cabral-Teixeira J, Selvendiran K, Kuppusamy P, Carvalho C, Beltran A, Moreno E, Pati UK, Gogna R. Nucleic Acids Res. 2019 Sep 20. pii: gkz766. doi: 10.1093/nar/gkz766. [Epub ahead of print] PMID: 31538203.

Times cited: 2

Altmetric score: 31


47. Madan E, Pelham CJ, Nagane M, Parker TM, Fazio K, Shaik K, Vieira A, Galzerano A, Canas-Marques R, Yuan Y, Yamashita T, Post SR, Soldini D, Yamashita H, Accardi D, Ferreira-Pinto MA, Camacho D, Hansen LA, Carvalho C, Beltran A, Kuppusamy P, Gogna R*, Moreno E*. Flower isoforms promote competitive growth in cancer. Nature. 2019 Aug;572(7768):260-264. doi: 10.1038/s41586-019-1429-3. Epub 2019 Jul 24. PMID: 31341286.

News and Views in Nature. Selected as one of the best cancer papers of 2019 by the EACR. Evaluated as “exceptional” in Faculty of 1000.

Times cited: 10

Altmetric score: 135


46. Coelho DS, Moreno E.

Emerging links between cell competition and Alzheimer’s disease. J Cell Sci. 2019 Jul 1;132(13). pii: jcs231258. doi: 10.1242/jcs.231258. Review. PMID: 31263078

Times cited:

Altmetric score: 3


45. 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. doi:10.1016/j.cub.2018.11.007. Epub 2018 Dec 13. PubMed PMID: 30554899.

Featured article in Current Biology. Editors’ choice in Science Signaling.

*Co-corresponding authors.

Times cited: 17

Altmetric score: 68


44. Smith JR et al. Proceedings of the Association for Research in Vision and Ophthalmology and Champalimaud Foundation Ocular Oncogenesis and Oncology Conference. Translational Vision Science & Technology. January 2019, Vol.8, 9. doi:10.1167/tvst.8.1.9. Perspective.

Times cited: 2

Altmetric score: 2


43. Coelho DS, Schwartz S, Merino MM, Hauert B, Topfel B, Tieche C, Rhiner C, Moreno E.

Culling Less Fit Neurons Protects against Amyloid-β-Induced Brain Damage and Cognitive and Motor Decline. CELL reports. 2018 Dec 26;25(13):3661-3673.e3. doi: 10.1016/j.celrep.2018.11.098. PMID: 30590040

Featured article in Cell reports.

Times cited: 4

Altmetric score: 119


42. Madan E, Gogna R, Moreno E.

Cell competition in development: information from flies and vertebrates. Curr Opin Cell Biol. 2018 Dec;55:150-157. doi:

10.1016/ Epub 2018 Sep 9. Review. PubMed PMID: 30208354.

Times cited: 11

Altmetric score: 8


41. *Germani, F., *Hain, D., Sternlicht, D., *Moreno, E., *Basler, K.

The Toll pathway inhibits tissue growth and regulates cell fitness in an infection-dependent manner. Elife. 2018 Nov 19;7. pii: e39939. doi: 10.7554/eLife.39939. PubMed PMID: 30451683; PubMed Central PMCID: PMC6279345.

*Co-corresponding authors, contributed equally to this work.

Times cited: 10

Altmetric score: 12


40. Brás-Pereira C, Moreno E.

Mechanical cell competition. Curr Opin Cell Biol.

2018 Apr;51:15-21. doi: 10.1016/ Epub 2017 Dec 6. Review.

PubMed PMID: 29153702.

Times cited: 15

Altmetric score: 8


39. Merino MM, Levayer R, Moreno E.

Survival of the Fittest: Essential Roles of Cell Competition in Development, Aging, and Cancer. Trends Cell Biol. 2016 Jun 16. pii: S0962-8924(16)30052-6. doi: 10.1016/j.tcb.2016.05.009.

Times cited: 65

Altmetric score: 8


38. Levayer R, Dupont C, Moreno E.

Tissue Crowding Induces Caspase-Dependent Competition for Space. Curr Biol. 2016 Feb 17. pii: S0960-9822(16)00059-2. doi: 10.1016/j.cub.2015.12.072. [Epub ahead of print]

PMID: 26898471

Times cited: 82

Altmetric score: 8


37. Levayer R, Moreno E.

How to be in a good shape? The influence of clone morphology on cell competition. Commun Integr Biol. 2016 Jan 5;9(1):e1102806. doi: 10.1080/19420889.2015.1102806. eCollection 2016 Jan-Feb. Review.

PMID: 27066183

Times cited: 9

Altmetric score: 2


36. Casas-Tinto, S., Lolo, F. and Moreno E.

Active JNK-dependent secretion of Drosophila Tyrosyl-tRNA synthetase by loser cells recruits haemocytes during cell competition. Nat. Commun. 6:10022 doi: 10.1038/ncomms10022 (2015).

Times cited: 23

Altmetric score: 12


35. Gogna R, Shee K, Moreno E.

Cell Competition During Growth and Regeneration. Annu. Rev. Genet. 2015 Nov 23;49:697-718. doi: 10.1146/annurev-genet-112414-055214.

PubMed PMID: 26631518.

Times cited: 26

Altmetric score: 9


34. Levayer, R., Hauert, B. and Moreno, E.

Cell mixing induced by myc is required for competitive tissue invasion and destruction. Nature. 2015 Aug 27; 524(7566):476-480. doi: 10.1038/nature14684. Epub 2015 Aug 19.

PMID: 26287461

Times cited: 68

Altmetric score: 48


33. Moreno E., Fernandez-Marrero, Y., Meyer, P., and Rhiner, C.

Brain regeneration in Drosophila involves comparison of neuronal fitness. Curr Biol. 2015 Mar 30;25(7):955-63. doi: 10.1016/j.cub.2015.02.014. Epub 2015 Mar 5.

PMID: 25754635

*Commentary in Current Biology.

Times cited: 19

Altmetric score: 7


32. Merino MM, Rhiner C, Lopez-Gay JM, Buechel D, Hauert B, Moreno E. Elimination of unfit cells maintains tissue health and prolongs lifespan. CELL. 2015 Jan29;160(3):461-76. doi: 10.1016/j.cell.2014.12.017. Epub 2015 Jan 15. PubMed PMID: 25601460.

*Commentaries in Current Biology, Washington Post, Daily Mail, NZZ, LabTimes, WIRED.

* Evaluated as “exceptional” in Faculty of 1000.

Times cited: 81

Altmetric score: 224


31.  Moreno, E. and Rhiner, C.

Darwin`s multicellularity: From neurotrophic theories and cell competition to fitness fingerprints. Current Opinion in Cell Biology, 31, 16-22. 2014. Doi: 10.1016/

Times cited: 25

Altmetric score: 5


30.  Moreno E.

Cancer: Darwinian tumour suppression. Nature. 2014 May 22;509(7501):435-6. doi: 10.1038/nature13337. Epub 2014 May 14. (News and Views).

Times cited: 11

Altmetric score: 8


29.  Zhang C, Li G, Casas-Tintó S, Lin N, Chung M, Moreno E, Moberg K and Zhou L. An Intergenic Regulatory Region Mediates Drosophila Myc -Induced Apoptosis and Blocks Tissue Hyperplasia. Oncogene. 2014 Jun 16. doi: 10.1038/onc.2014.160.

Times cited: 12

Altmetric score: 1


28. K Gupta, E Madan, M Sayyid, H Arias-Pulido, E Moreno, P Kuppusamy, R Gogna. Oxygen regulates molecular mechanisms of cancer progression and metastasis. Cancer and Metastasis Reviews, Volume 33, Issue 1 , pp 183-215. 2014.

Times cited: 6


27. Merino MM, Rhiner C, Portela M, Moreno E.

“Fitness fingerprints” mediate physiological culling of unwanted neurons in Drosophila. Curr Biol. 2013 Jul 22;23(14):1300-9. doi: 10.1016/j.cub.2013.05.053. Epub 2013 Jun 27.

Times cited: 38

Altmetric score: 13

*Commentary in Current Biology.


26. Fernandez-Hernandez, I., Rhiner, C., and Moreno, E. (2013).

Adult Neurogenesis in Drosophila. CELL reports, Jun 27;3(6):1857-65. doi: 10.1016/j.celrep.2013.05.034. Epub 2013 Jun 20.

Times cited: 63

Altmetric score: 20 

* Evaluated as “exceptional” in Faculty of 1000. Selected among best papers published in Cell Reports in 2013.


25. Moreno E.

The “Out of Africa Tribe” (II): Paleolithic warriors with big canoes and protective weapons. Commun Integr Biol. 2013 May 1;6(3):e24145. doi:


Times cited: 6

Altmetric score: 2


24. Lolo, F., Casas-Tinto S. and Moreno E.

How winner cells cause the demise of loser cells. Bioessays, 2013 (AOP). DOI: 10.1002/bies.201200156

Times cited: 8

Altmetric score: 4

* Featured in Medfriendly.


23. Levayer, R. and Moreno, E.

Mechanisms of cell competition: Themes and variations. Journal of Cell Biology (JCB), 2013 Mar 18;200(6):689-98. doi: 10.1083/jcb.201301051.

Times cited: 114

Altmetric score: 4


22. Lolo, F., Casas-Tinto S. and Moreno E.

Cell competition timeline: winners kill losers, which are extruded and engulfed by hemocytes. CELL reports, 2 (2012) pp. 526-539. DOI: 10.1016/j.celrep.2012.08.012.

Times cited: 63

Altmetric score: 31 


21. Moreno E.

Design and Construction of ‘‘Synthetic Species’’. PLoS ONE (2012) 7(7): e39054. doi:10.1371/journal.pone.0039054.

Times cited: 16

Altmetric score: 122

* PlosOne editor´s choice in Plos blogs. Featured in LabTimes and Greenwire. Selected as one of the “100 discoveries of the year 2012” by “Quarks and Co”.


20. Petrova E, López-Gay JM, Rhiner C, Moreno E.

Flower-deficient mice have reduced susceptibility to skin papilloma formation. Dis Model Mech. 2012 Jul;5(4):553-61.

Times Cited: 32

Altmetric score: 2

* Commentary in Disease Models and Mechanisms.


19. Staal FJ et al. Stem cell self-renewal: lessons from bone marrow, gut and iPS toward clinical applications. Leukemia. 2011 Jul;25(7):1095-102. doi:10.1038/leu.2011.52. Epub 2011 Apr 29. Review. PubMed PMID: 21527933.

Times cited: 29


18. Petrova E, Soldini D and Moreno E.

The expression of SPARC in human tumors is consistent with its role during cell competition. Communicative and Integrative Biology. Volume 4, Issue 2 March/April 2011, 171 – 174.

Times Cited: 24


17. Moreno E. The society of our “out of Africa” ancestors (I): The migrant warriors that colonized the world. Communicative and Integrative Biology. Vol. 4, Issue 2   March/April 2011 163 – 170.

Times Cited: 25 (including pre-print citations).

Altmetric score: 2


16. Casas-Tinto S, Torres M and Moreno E.

The “Flower Code” and Cancer Development.

Clin Transl Oncol. 2011 Jan;13(1):5-9.

Times cited: 9


15. Portela M, Casas-Tinto S, Rhiner C, López-Gay JM, Domínguez O, Soldini D, Moreno E.

Drosophila SPARC is a self-protective signal expressed by loser cells during cell competition. DEV. CELL. 2010. Vol. 19, Issue Issue 4, pp. 562-573.

Times Cited: 96

Altmetric score: 3

* Evaluated as “very good” in Faculty of 1000.

*Commentaries in The Scientist and Nature Reviews Cancer.


14. Rhiner C, López-Gay JM, Soldini D, Casas-Tinto S, Martín FA, Lombardía L, Moreno E.

Flower Forms an Extracellular Code that Reveals the Fitness of a Cell to its Neighbors in Drosophila. DEV. CELL. Vol. 18, Issue 6, 985-998, 2010.

Times Cited: 130

Altmetric score: 11

* Evaluated as “very good” in Faculty of 1000.

*Commentaries in Nature Reviews Genetics and Developmental Cell.


13. Ziv O, Suissa Y, Neuman H, Dinur T, Geuking P, Rhiner C, Portela M, Lolo F, Moreno E*, and Gerlitz O*.

The co-regulator dNAB interacts with Brinker to eliminate cells with reduced Dpp signalling. DEVELOPMENT. 2009. 136: 1137-1145.

* Co-corresponding authors.

Times Cited: 22


12. Rhiner C, Díaz B, Portela M, Poyatos JF, Fernández-Ruiz I, López-Gay JM, Gerlitz O, Moreno E.

Persistent competition among stem cells and their daughters in the Drosophila ovary germline niche. DEVELOPMENT. 2009. 136: 995-1006  

Times Cited: 71

Altmetric score: 2

* Evaluated as “very good” in Faculty of 1000.


11. Rhiner C, Moreno E.

Super competition as a possible mechanism to pioneer pre-cancerous fields. CARCINOGENESIS. 2009 Jan 6. 30(5): 723-728.

Times Cited: 52


10. Soldini D*, Moreno E*, Martin V, Gratwohl A, Marone C, Mazzucchelli L.

BM-derived cells randomly contribute to neoplastic and non-neoplastic epithelial tissues at low rates. BONE MARROW TRANSPLANT. 2008 Dec;42(11):749-55.

Times Cited: 9

* Equal contribution.


9. Gerlitz O, Wagner EF, Moreno E.

When flies and mice develop cancer. Meeting on development and cancer. EMBO Rep. 2008 Aug;9(8):730-734.


8. Moreno E.

Is cell competition relevant to cancer? NAT REV CANCER. FEB; 8(2):141-7, 2008.

Times Cited: 166

Altmetric score: 6


7. Diaz B, Moreno E

The competitive nature of cells. EXPERIMENTAL CELL RESEARCH.

306 (2): 317-322 JUN 10, 2005.

Times Cited: 56


6. Martin FA, Perez-Garijo A, Moreno E, Morata G.

The brinker gradient controls wing growth in Drosophila

DEVELOPMENT 131 (20): 4921-4930 OCT, 2004.

Times Cited: 107


5. Moreno E, Basler K

dMyc transforms cells into super-competitors

CELL 117 (1): 117-129 APR 2, 2004.

Times Cited: 469

Altmetric score: 1

News and Views in Nature.

* Evaluated as “exceptional” in Faculty of 1000.


4. Moreno E, Yan MH, Basler K

Evolution of TNF signaling mechanisms: JNK-dependent apoptosis triggered by Eiger, the Drosophila homolog of the TNF superfamily

CURRENT BIOLOGY 12 (14): 1263-1268 JUL 23, 2002.

Times Cited: 308

Altmetric score: 2

* Evaluated as “very good” in Faculty of 1000.


3. Moreno E, Basler K, Morata G

Cells compete for Decapentaplegic survival factor to prevent apoptosis in Drosophila wing development

NATURE 416 (6882): 755-759 APR 18, 2002.

Times Cited: 364

Altmetric score: 1

Evaluated 5 in Faculty of 1000.


2. Moreno E, Morata G

Caudal is the Hox gene that specifies the most posterior Drosophila segment

NATURE 400 (6747): 873-877 AUG 26, 1999.

Times Cited: 128


1. Calleja M, Moreno E, Pelaz S, Morata G.

Visualization of gene expression in living adult Drosophila

SCIENCE 274 (5285): 252-255 OCT 11, 1996.

Times Cited: 457