J. Alvar, I. Vélez, C. Bern, M. Herrero, and P. Desjeux, Leishmaniasis Worldwide and Global Estimates of Its Incidence, PLoS ONE, vol.83, issue.30, p.35671, 2012.
DOI : 10.1371/journal.pone.0035671.s101

D. Zilberstein and M. Shapira, The Role of pH and Temperature in the Development of Leishmania Parasites, Annual Review of Microbiology, vol.48, issue.1, pp.449-470, 1994.
DOI : 10.1146/annurev.mi.48.100194.002313

P. Doyle, J. Engel, P. Pimenta, P. Da-silva, and D. Dwyer, Leishmania donovani: Long-term culture of axenic amastigotes at 37 ??C, Experimental Parasitology, vol.73, issue.3, pp.326-334, 1991.
DOI : 10.1016/0014-4894(91)90104-5

S. Goyard, H. Segawa, J. Gordon, M. Showalter, and R. Duncan, An in vitro system for developmental and genetic studies of Leishmania donovani phosphoglycans, Molecular and Biochemical Parasitology, vol.130, issue.1, pp.31-42, 2003.
DOI : 10.1016/S0166-6851(03)00142-7

D. Kültz, Phylogenetic and Functional Classification of Mitogen- and Stress-Activated Protein Kinases, Journal of Molecular Evolution, vol.46, issue.5, pp.571-588, 1998.
DOI : 10.1007/PL00006338

A. Hindley and W. Kolch, Extracellular signal regulated kinase (ERK)/ mitogen activated protein kinase (MAPK)-independent functions of Raf kinases, J Cell Sci, vol.115, pp.1575-1581, 2002.

J. Ferrell, J. Bhatt, and R. , Mechanistic Studies of the Dual Phosphorylation of Mitogen-activated Protein Kinase, Journal of Biological Chemistry, vol.272, issue.30, pp.19008-19016, 1997.
DOI : 10.1074/jbc.272.30.19008

R. Davis, Transcriptional regulation by MAP kinases, Molecular Reproduction and Development, vol.367, issue.4, pp.459-467, 1995.
DOI : 10.1002/mrd.1080420414

M. Cargnello and P. Roux, Activation and Function of the MAPKs and Their Substrates, the MAPK-Activated Protein Kinases, Microbiology and Molecular Biology Reviews, vol.75, issue.1, pp.50-83, 2011.
DOI : 10.1128/MMBR.00031-10

M. Cheng, V. Sexl, C. Sherr, and M. Roussel, Assembly of cyclin D-dependent kinase and titration of p27Kip1 regulated by mitogen-activated protein kinase kinase (MEK1), Proceedings of the National Academy of Sciences, vol.95, issue.3, pp.1091-1096, 1998.
DOI : 10.1073/pnas.95.3.1091

J. Wright, E. Munar, D. Jameson, P. Andreassen, and R. Margolis, Mitogen-activated protein kinase kinase activity is required for the G2/M transition of the cell cycle in mammalian fibroblasts, Proceedings of the National Academy of Sciences, vol.96, issue.20, pp.11335-11340, 1999.
DOI : 10.1073/pnas.96.20.11335

D. Brooks, R. Mcculloch, G. Coombs, and J. Mottram, Stable transformation of trypanosomatids through targeted chromosomal integration of the selectable marker gene encoding blasticidin S deaminase, FEMS Microbiology Letters, vol.186, issue.2, pp.287-291, 2000.
DOI : 10.1111/j.1574-6968.2000.tb09119.x

J. Dean, M. Brook, A. Clark, and J. Saklatvala, p38 Mitogen-activated Protein Kinase Regulates Cyclooxygenase-2 mRNA Stability and Transcription in Lipopolysaccharide-treated Human Monocytes, Journal of Biological Chemistry, vol.274, issue.1, pp.264-269, 1999.
DOI : 10.1074/jbc.274.1.264

D. Parry, D. Mahony, K. Wills, and E. Lees, Cyclin D-CDK Subunit Arrangement Is Dependent on the Availability of Competing INK4 and p21 Class Inhibitors, Molecular and Cellular Biology, vol.19, issue.3, pp.1775-1783, 1999.
DOI : 10.1128/MCB.19.3.1775

A. Ivens, C. Peacock, E. Worthey, L. Murphy, and G. Aggarwal, The Genome of the Kinetoplastid Parasite, Leishmania major, Science, vol.309, issue.5733, pp.436-442, 2005.
DOI : 10.1126/science.1112680

M. Wiese, Leishmania MAP kinases ??? Familiar proteins in an unusual context, International Journal for Parasitology, vol.37, issue.10, pp.1053-1062, 2007.
DOI : 10.1016/j.ijpara.2007.04.008

M. Parsons, E. Worthey, P. Ward, and J. Mottram, Comparative analysis of the kinomes of three pathogenic trypanosomatids: Leishmania major, Trypanosoma brucei and Trypanosoma cruzi, BMC Genomics, vol.6, issue.1, p.127, 2005.
DOI : 10.1186/1471-2164-6-127

M. Wiese, A mitogen-activated protein (MAP) kinase homologue of Leishmania mexicana is essential for parasite survival in the infected host, The EMBO Journal, vol.17, issue.9, pp.2619-2628, 1998.
DOI : 10.1093/emboj/17.9.2619

M. Wiese, D. Kuhn, and C. Grünfelder, Protein Kinase Involved in Flagellar-Length Control, Eukaryotic Cell, vol.2, issue.4, pp.769-777, 2003.
DOI : 10.1128/EC.2.4.769-777.2003

F. Bengs, A. Scholz, D. Kuhn, and M. Wiese, LmxMPK9, a mitogen-activated protein kinase homologue affects flagellar length in Leishmania mexicana, Molecular Microbiology, vol.277, issue.5, pp.1606-1615, 2005.
DOI : 10.1111/j.1365-2958.2005.04498.x

D. Kuhn and M. Wiese, LmxPK4, a mitogen-activated protein kinase kinase homologue of Leishmania mexicana with a potential role in parasite differentiation, Molecular Microbiology, vol.48, issue.5, pp.1169-1182, 2005.
DOI : 10.1111/j.1365-2958.2005.04614.x

M. Erdmann, A. Scholz, I. Melzer, C. Schmetz, and M. Wiese, Interacting Protein Kinases Involved in the Regulation of Flagellar Length, Molecular Biology of the Cell, vol.17, issue.4, pp.2035-2045, 2006.
DOI : 10.1091/mbc.E05-10-0976

B. Rotureau, M. Morales, P. Bastin, and G. Spä-th, The flagellum-mitogen-activated protein kinase connection in Trypanosomatids: a key sensory role in parasite signalling and development?, Cellular Microbiology, vol.48, issue.5, pp.710-718, 2009.
DOI : 10.1111/j.1462-5822.2009.01295.x

M. Morales, O. Renaud, W. Faigle, S. Shorte, and G. Spä-th, Over-expression of Leishmania major MAP kinases reveals stage-specific induction of phosphotransferase activity, International Journal for Parasitology, vol.37, issue.11, pp.1187-1199, 2007.
DOI : 10.1016/j.ijpara.2007.03.006

M. Morales, R. Watanabe, C. Laurent, P. Lenormand, and J. Rousselle, Phosphoproteomic analysis ofLeishmania donovani pro- and amastigote stages, PROTEOMICS, vol.38, issue.2, pp.350-363, 2008.
DOI : 10.1002/pmic.200700697

M. Morales, P. Pescher, and G. Spä-th, Leishmania major MPK7 Protein Kinase Activity Inhibits Intracellular Growth of the Pathogenic Amastigote Stage, Eukaryotic Cell, vol.9, issue.1, pp.22-30, 2010.
DOI : 10.1128/EC.00196-09

URL : https://hal.archives-ouvertes.fr/pasteur-01433564

S. Von-freyend, H. Rosenqvist, A. Fink, I. Melzer, and J. Clos, LmxMPK4, an essential mitogen-activated protein kinase of Leishmania mexicana is phosphorylated and activated by the STE7-like protein kinase LmxMKK5, International Journal for Parasitology, vol.40, issue.8, pp.969-978, 2010.
DOI : 10.1016/j.ijpara.2010.02.004

N. Kannan, A. Neuwald, . Mapk, . Cdk, . Gsk et al., Evolutionary constraints associated with functional specificity of the CMGC protein kinases MAPK, CDK, GSK, SRPK, DYRK, and CK2??, Protein Science, vol.56, issue.8, pp.2059-2077, 2004.
DOI : 10.1110/ps.04637904

S. Horjales, D. Schmidt-arras, R. Limardo, O. Leclercq, and G. Obal, The Crystal Structure of the MAP Kinase LmaMPK10 from Leishmania Major Reveals Parasite-Specific Features and Regulatory Mechanisms, Structure, vol.20, issue.10, pp.1649-1660, 2012.
DOI : 10.1016/j.str.2012.07.005

URL : https://hal.archives-ouvertes.fr/pasteur-00727033

G. Zhou, Z. Bao, and J. Dixon, Components of a new human protein kinase signal transduction pathway, J Biol Chem, vol.270, pp.12665-12669, 1995.

J. Lee, R. Ulevitch, and J. Han, Primary Structure of BMK1: A New Mammalian MAP Kinase, Biochemical and Biophysical Research Communications, vol.213, issue.2, pp.715-724, 1995.
DOI : 10.1006/bbrc.1995.2189

J. Ellis, M. Sarkar, E. Hendriks, and K. Matthews, A novel ERK-like, CRK-like protein kinase that modulates growth in Trypanosoma brucei via an autoregulatory C-terminal extension, Molecular Microbiology, vol.254, issue.5, pp.1487-1499, 2004.
DOI : 10.1111/j.1365-2958.2004.04218.x

M. Abe, W. Kuo, M. Hershenson, and M. Rosner, Extracellular Signal-Regulated Kinase 7 (ERK7), a Novel ERK with a C-Terminal Domain That Regulates Its Activity, Its Cellular Localization, and Cell Growth, Molecular and Cellular Biology, vol.19, issue.2, pp.1301-1312, 1999.
DOI : 10.1128/MCB.19.2.1301

M. Abe, K. Kahle, M. Saelzler, K. Orth, and J. Dixon, ERK7 Is an Autoactivated Member of the MAPK Family, Journal of Biological Chemistry, vol.276, issue.24, pp.21272-21279, 2001.
DOI : 10.1074/jbc.M100026200

A. Kornev, N. Haste, S. Taylor, and L. Eyck, Surface comparison of active and inactive protein kinases identifies a conserved activation mechanism, Proceedings of the National Academy of Sciences, vol.103, issue.47, pp.17783-17788, 2006.
DOI : 10.1073/pnas.0607656103

S. Taylor and A. Kornev, Protein kinases: evolution of dynamic regulatory proteins, Trends in Biochemical Sciences, vol.36, issue.2, pp.65-77, 2011.
DOI : 10.1016/j.tibs.2010.09.006

M. Cobb, T. Boulton, and D. Robbins, Extracellular signal-regulated kinases: ERKs in progress., Molecular Biology of the Cell, vol.2, issue.12, pp.965-978, 1991.
DOI : 10.1091/mbc.2.12.965

M. Verlhac, J. Kubiak, H. Clarke, and B. Maro, Microtubule and chromatin behavior follow MAP kinase activity but not MPF activity during meiosis in mouse oocytes, Development, vol.120, pp.1017-1025, 1994.

J. Kubiak, Protein Kinase Assays for Measuring MPF and MAPK Activities in Mouse and Rat Oocytes and Early Embryos, Methods Mol Biol, vol.957, pp.77-89, 2013.
DOI : 10.1007/978-1-62703-191-2_5

URL : https://hal.archives-ouvertes.fr/inserm-00633900

H. Rosenqvist, J. Ye, and O. Jensen, Analytical Strategies in Mass Spectrometry-Based Phosphoproteomics, Methods Mol Biol, vol.753, pp.183-213, 2011.
DOI : 10.1007/978-1-61779-148-2_13

C. Prowse and J. Lew, Mechanism of Activation of ERK2 by Dual Phosphorylation, Journal of Biological Chemistry, vol.276, issue.1, pp.99-103, 2001.
DOI : 10.1074/jbc.M008137200

D. Robbins, E. Zhen, H. Owaki, C. Vanderbilt, and D. Ebert, Regulation and properties of extracellular signal-regulated protein kinases 1 and 2 in vitro, J Biol Chem, vol.268, pp.5097-5106, 1993.

M. Cobb and E. Goldsmith, How MAP Kinases Are Regulated, Journal of Biological Chemistry, vol.270, issue.25, pp.14843-14846, 1995.
DOI : 10.1074/jbc.270.25.14843

M. Morales, R. Watanabe, M. Dacher, P. Chafey, and J. Osorio-y-fortéa, Phosphoproteome dynamics reveal heat-shock protein complexes specific to the Leishmania donovani infectious stage, Proceedings of the National Academy of Sciences, vol.107, issue.18, pp.8381-8386, 2010.
DOI : 10.1073/pnas.0914768107

URL : https://hal.archives-ouvertes.fr/pasteur-00476914

M. Huse and J. Kuriyan, The Conformational Plasticity of Protein Kinases, Cell, vol.109, issue.3, pp.275-282, 2002.
DOI : 10.1016/S0092-8674(02)00741-9

B. Canagarajah, A. Khokhlatchev, M. Cobb, and E. Goldsmith, Activation Mechanism of the MAP Kinase ERK2 by Dual Phosphorylation, Cell, vol.90, issue.5, pp.859-869, 1997.
DOI : 10.1016/S0092-8674(00)80351-7

M. Bell and D. Engelberg, Phosphorylation of Tyr-176 of the Yeast MAPK Hog1/p38 Is Not Vital for Hog1 Biological Activity, Journal of Biological Chemistry, vol.278, issue.17, pp.14603-14606, 2003.
DOI : 10.1074/jbc.C300006200

L. Johnson, M. Noble, and D. Owen, Active and Inactive Protein Kinases: Structural Basis for Regulation, Cell, vol.85, issue.2, pp.149-158, 1996.
DOI : 10.1016/S0092-8674(00)81092-2

J. English, G. Pearson, J. Wilsbacher, J. Swantek, and M. Karandikar, New Insights into the Control of MAP Kinase Pathways, Experimental Cell Research, vol.253, issue.1, pp.255-270, 1999.
DOI : 10.1006/excr.1999.4687

J. Gonzalez, A. Cornejo, M. Santos, E. Cordero, and B. Gutierrez, A novel protein phosphatase 2A (PP2A) is involved in the transformation of human protozoan parasite Trypanosoma cruzi, Biochemical Journal, vol.374, issue.3, pp.647-656, 2003.
DOI : 10.1042/bj20030215

M. Buschbeck and A. Ullrich, The Unique C-terminal Tail of the Mitogen-activated Protein Kinase ERK5 Regulates Its Activation and Nuclear Shuttling, Journal of Biological Chemistry, vol.280, issue.4, pp.2659-2667, 2005.
DOI : 10.1074/jbc.M412599200

S. Hu, M. Parker, J. Lei, M. Wilce, and G. Benian, Insights into autoregulation from the crystal structure of twitchin kinase, Nature, vol.369, issue.6481, pp.581-584, 1994.
DOI : 10.1038/369581a0

J. Brondello, J. Pouyssegur, and F. Mckenzie, Reduced MAP Kinase Phosphatase-1 Degradation After p42/p44MAPK-Dependent Phosphorylation, Science, vol.286, issue.5449, pp.2514-2517, 1999.
DOI : 10.1126/science.286.5449.2514

A. Nichols, M. Camps, C. Gillieron, C. Chabert, and A. Brunet, Substrate recognition domains within extracellular signal-regulated kinase mediate binding and catalytic activation of mitogen, 2000.

T. Smith, D. Sweetman, M. Patterson, S. Keyse, and A. Munsterberg, Feedback interactions between MKP3 and ERK MAP kinase control scleraxis expression and the specification of rib progenitors in the developing chick somite, Development, vol.132, issue.6, pp.1305-1314, 2005.
DOI : 10.1242/dev.01699

W. Breitwieser, S. Lyons, A. Flenniken, G. Ashton, and G. Bruder, Feedback regulation of p38 activity via ATF2 is essential for survival of embryonic liver cells, Genes & Development, vol.21, issue.16, pp.2069-2082, 2007.
DOI : 10.1101/gad.430207

G. Cohen-freue, T. Holzer, J. Forney, and W. Mcmaster, Global gene expression in Leishmania, International Journal for Parasitology, vol.37, issue.10, pp.1077-1086, 2007.
DOI : 10.1016/j.ijpara.2007.04.011

N. Mody, D. Campbell, N. Morrice, P. M. Cohen, and P. , An analysis of the phosphorylation and activation of extracellular-signal-regulated protein kinase 5 (ERK5) by mitogen-activated protein kinase kinase 5 (MKK5) in vitro, Biochemical Journal, vol.372, issue.2, pp.567-75, 2003.
DOI : 10.1042/bj20030193

Y. Saar, A. Ransford, E. Waldman, S. Mazareb, and S. Amin-spector, Characterization of developmentally-regulated activities in axenic amastigotes of Leishmania donovani, Molecular and Biochemical Parasitology, vol.95, issue.1, pp.9-20, 1998.
DOI : 10.1016/S0166-6851(98)00062-0

D. Dwyer, Antibody-induced modulation of Leishmania donovani surface membrane antigens, J Immunol, vol.117, pp.2081-2091, 1976.

B. Menz, G. Winter, T. Ilg, F. Lottspeich, and P. Overath, Purification and characterization of a membrane-bound acid phosphatase of Leishmania mexicana, Molecular and Biochemical Parasitology, vol.47, issue.1, pp.101-108, 1991.
DOI : 10.1016/0166-6851(91)90152-V

J. Lebowitz, Chapter 4: Transfection Experiments with Leishmania, Methods Cell Biol, vol.45, pp.65-78, 1994.
DOI : 10.1016/S0091-679X(08)61846-4

M. Aslett, C. Aurrecoechea, M. Berriman, J. Brestelli, and B. Brunk, TriTrypDB: a functional genomic resource for the Trypanosomatidae, Nucleic Acids Research, vol.38, issue.Database, pp.457-462, 2010.
DOI : 10.1093/nar/gkp851

S. Altschul, T. Madden, A. Schä-ffer, J. Zhang, and Z. Zhang, Gapped BLAST and PSI-BLAST: a new generation of protein database search programs, Nucleic Acids Research, vol.25, issue.17, pp.3389-3402, 1997.
DOI : 10.1093/nar/25.17.3389

I. Jonassen, Efficient discovery of conserved patterns using a pattern graph, Bioinformatics, vol.13, issue.5, pp.509-522, 1997.
DOI : 10.1093/bioinformatics/13.5.509