P. Ananatis-grazing-resistance-bergman, K. Arends, J. Schölvinck, and E. , Issue 1 e00075-15 msphere.asm.org 13 REFERENCES 1 Pantoea agglomerans septicemia in three newborn infants, Pediatr Infect Dis J, vol.1, issue.26, pp.453-454, 2007.

A. Shubov, P. Jagannathan, and P. Chin-hong, Pantoea agglomerans pneumonia in a heart-lung transplant recipient: case report and a review of an emerging pathogen in immunocompromised hosts, Transplant Infectious Disease, vol.300, issue.(16), pp.536-539, 2011.
DOI : 10.1111/j.1399-3062.2011.00630.x

K. Lalas and D. Erichsen, Sporadic Pantoea agglomerans bacteremia in a near-term female: case report and review of literature, Jpn J Infect Dis, vol.63, pp.290-291, 2010.

A. Cheng, C. Liu, H. Tsai, M. Hsu, C. Yang et al., Bacteremia caused by Pantoea agglomerans at a medical center in Taiwan, 2000???2010, Journal of Microbiology, Immunology and Infection, vol.46, issue.3, pp.187-194, 2000.
DOI : 10.1016/j.jmii.2012.05.005

A. Cruz, A. Cazacu, and C. Allen, Pantoea agglomerans, a Plant Pathogen Causing Human Disease, Journal of Clinical Microbiology, vol.45, issue.6, pp.632-639, 1989.
DOI : 10.1128/JCM.00632-07

D. Champs, C. , L. Seaux, S. Dubost, J. Boisgard et al., Isolation of Pantoea agglomerans in two cases of septic monoarthritis after plant thorn and wood sliver injuries, J Clin Microbiol, vol.38, pp.460-461, 2000.

H. Habsah, M. Zeehaida, H. Van-rostenberghe, R. Noraida, W. Pauzi et al., An outbreak of Pantoea spp. in a neonatal intensive care unit secondary to contaminated parenteral nutrition, Journal of Hospital Infection, vol.61, issue.3, pp.213-218, 2005.
DOI : 10.1016/j.jhin.2005.01.004

T. Coutinho and S. Venter, : an unconventional plant pathogen, Molecular Plant Pathology, vol.40, issue.3, pp.325-335, 2009.
DOI : 10.1111/j.1364-3703.2009.00542.x

A. Walterson and J. Stavrinides, insights into a highly versatile and diverse genus within the Enterobacteriaceae, FEMS Microbiology Reviews, vol.39, issue.6, pp.968-984, 2015.
DOI : 10.1093/femsre/fuv027

F. Serrano, Bacterial fruitlet brown rot on pineapple in the Philippines, Philipp J Sci, vol.36, pp.271-305, 1928.

B. Völksch, S. Thon, I. Jacobsen, and M. Gube, Polyphasic study of plant- and clinic-associated Pantoea agglomerans strains reveals indistinguishable virulence potential, Infection, Genetics and Evolution, vol.9, issue.6, pp.1381-1391, 2009.
DOI : 10.1016/j.meegid.2009.09.016

G. Nadarasah and J. Stavrinides, Quantitative evaluation of the host-colonizing capabilities of the enteric bacterium Pantoea using plant and insect hosts, Microbiology, vol.160, issue.Pt_3, pp.602-615, 2014.
DOI : 10.1099/mic.0.073452-0

M. Kirzinger, G. Nadarasah, and J. Stavrinides, Insights into crosskingdom plant pathogenic bacteria, Genes (Basel), vol.2, pp.980-997, 2011.

G. Nadarasah and J. Stavrinides, Insects as alternative hosts for phytopathogenic bacteria, FEMS Microbiology Reviews, vol.35, issue.3, pp.555-575, 2011.
DOI : 10.1111/j.1574-6976.2011.00264.x

J. Stavrinides, A. No, and H. Ochman, enables gut colonization and pathogenicity in an insect host, Environmental Microbiology, vol.175, issue.1, pp.147-155, 2010.
DOI : 10.1111/j.1462-2920.2009.02056.x

D. Maayer, P. Chan, W. Rubagotti, E. Venter, S. Toth et al., Analysis of the Pantoea ananatis pan-genome reveals factors underlying its ability to colonize and interact with plant, insect and vertebrate hosts, BMC Genomics, vol.15, issue.1, pp.4041471-2164, 2014.
DOI : 10.1128/JB.187.18.6258-6264.2005

D. Shyntum, S. Venter, L. Moleleki, I. Toth, and T. Coutinho, Comparative genomics of type VI secretion systems in strains of Pantoea ananatis from different environments, BMC Genomics, vol.15, issue.1, p.163, 2014.
DOI : 10.1093/nar/28.1.33

M. Kirzinger and J. Stavrinides, Host specificity determinants as a genetic continuum, Trends in Microbiology, vol.20, issue.2, pp.88-93, 2012.
DOI : 10.1016/j.tim.2011.11.006

M. Kirzinger, C. Butz, and J. Stavrinides, Inheritance of Pantoea type III secretion systems through both vertical and horizontal transfer, Molecular Genetics and Genomics, vol.9, issue.5, 2015.
DOI : 10.1007/s00438-015-1062-2

P. Cosson and T. Soldati, Eat, kill or die: when amoeba meets bacteria, Current Opinion in Microbiology, vol.11, issue.3, pp.271-276, 2008.
DOI : 10.1016/j.mib.2008.05.005

B. Hasselbring, M. Patel, and M. Schell, Dictyostelium discoideum as a model system for identification of Burkholderia pseudomallei viru- Smith et al, 2011.

Y. Pan, T. Lin, C. Hsu, and J. Wang, Use of a Dictyostelium Model for Isolation of Genetic Loci Associated with Phagocytosis and Virulence in Klebsiella pneumoniae, Infection and Immunity, vol.79, issue.3, pp.997-1006, 2011.
DOI : 10.1128/IAI.00906-10

E. Lelong, A. Marchetti, M. Simon, J. Burns, C. Van-delden et al., Evolution of Pseudomonas aeruginosa virulence in infected patients revealed in a Dictyostelium discoideum host model, Clinical Microbiology and Infection, vol.17, issue.9, pp.1415-1420, 2011.
DOI : 10.1111/j.1469-0691.2010.03431.x

L. Alibaud, T. Köhler, A. Coudray, C. Prigent-combaret, E. Bergeret et al., Pseudomonas aeruginosa virulence genes identified in a Dictyostelium host model, Cellular Microbiology, vol.179, issue.11, pp.729-740, 2008.
DOI : 10.1128/IAI.01772-05

URL : https://hal.archives-ouvertes.fr/hal-00338552

R. Froquet, E. Lelong, A. Marchetti, and P. Cosson, Dictyostelium discoideum: a model host to measure bacterial virulence, Nature Protocols, vol.275, issue.1, pp.25-30, 2008.
DOI : 10.1038/nprot.2008.212

S. Pukatzki, R. Kessin, and J. Mekalanos, The human pathogen Pseudomonas aeruginosa utilizes conserved virulence pathways to infect the social amoeba Dictyostelium discoideum, Proceedings of the National Academy of Sciences, vol.99, issue.5, pp.3159-3164, 2002.
DOI : 10.1073/pnas.052704399

D. Macintyre, S. Miyata, M. Kitaoka, and S. Pukatzki, The Vibrio cholerae type VI secretion system displays antimicrobial properties, Proceedings of the National Academy of Sciences, vol.107, issue.45, pp.19520-19524, 2010.
DOI : 10.1073/pnas.1012931107

D. Sperandio, V. Decoin, X. Latour, L. Mijouin, M. Hillion et al., Virulence of the Pseudomonas fluorescens clinical strain MFN1032 towards Dictyostelium discoideum and macrophages in relation with type III secretion system, BMC Microbiology, vol.12, issue.1, p.223, 2012.
DOI : 10.1186/1471-2180-12-223

M. Totsika, B. Heras, D. Wurpel, and M. Schembri, Characterization of Two Homologous Disulfide Bond Systems Involved in Virulence Factor Biogenesis in Uropathogenic Escherichia coli CFT073, Journal of Bacteriology, vol.191, issue.12, pp.3901-390800143, 2009.
DOI : 10.1128/JB.00143-09

A. Qin, D. Scott, and B. Mann, Francisella tularensis subsp. tularensis Schu S4 Disulfide Bond Formation Protein B, but Not an RND-Type Efflux Pump, Is Required for Virulence, Infection and Immunity, vol.76, issue.7, pp.3086-3092, 2008.
DOI : 10.1128/IAI.00363-08

K. Denoncin and J. Collet, Disulfide Bond Formation in the Bacterial Periplasm: Major Achievements and Challenges Ahead, Antioxidants & Redox Signaling, vol.19, issue.1, pp.63-71, 2013.
DOI : 10.1089/ars.2012.4864

E. Bjur, S. Eriksson-ygberg, F. Åslund, and M. Rhen, Thioredoxin 1 Promotes Intracellular Replication and Virulence of Salmonella enterica Serovar Typhimurium, Infection and Immunity, vol.74, issue.9, pp.5140-5151, 2006.
DOI : 10.1128/IAI.00449-06

A. Abdel-mawgoud, R. Hausmann, F. Lépine, M. Müller, and E. Déziel, Rhamnolipids: Detection, Analysis, Biosynthesis, Genetic Regulation, and Bioengineering of Production, Biosurfactants: from genes to applications, pp.13-55, 2011.
DOI : 10.1007/978-3-642-14490-5_2

M. Alexeyev, I. Shokolenko, and T. Croughan, Improved antibiotic-resistance gene cassettes and omega elements for Escherichia coli vector construction and in vitro deletion/insertion mutagenesis, Gene, vol.160, issue.1, 1995.
DOI : 10.1016/0378-1119(95)00108-I

M. Alexeyev, I. Shokolenko, and T. Croughan, derivatives for insertion mutagenesis and genetic engineering in Gram-negative bacteria, Canadian Journal of Microbiology, vol.41, issue.11, pp.1053-1055, 1995.
DOI : 10.1139/m95-147

A. Abdel-mawgoud, F. Lépine, and E. Déziel, Rhamnolipids: diversity of structures, microbial origins and roles, Applied Microbiology and Biotechnology, vol.74, issue.13, pp.1323-1336, 2010.
DOI : 10.1007/s00253-010-2498-2

URL : https://hal.archives-ouvertes.fr/pasteur-00819624

N. Caiazza, R. Shanks, O. Toole, and G. , Rhamnolipids Modulate Swarming Motility Patterns of Pseudomonas aeruginosa, Journal of Bacteriology, vol.187, issue.21, pp.7351-7361, 2005.
DOI : 10.1128/JB.187.21.7351-7361.2005

E. Déziel, F. Lépine, S. Milot, and R. Villemur, rhlA is required for the production of a novel biosurfactant promoting swarming motility in Pseudomonas aeruginosa: 3-(3-hydroxyalkanoyloxy)alkanoic acids (HAAs), the precursors of rhamnolipids, Microbiology, vol.149, issue.8, pp.2005-2013, 2003.
DOI : 10.1099/mic.0.26154-0

A. Nickzad, F. Lépine, and E. Déziel, Quorum Sensing Controls Swarming Motility of Burkholderia glumae through Regulation of Rhamnolipids, PLOS ONE, vol.79, issue.1, 2015.
DOI : 10.1371/journal.pone.0128509.t002

URL : https://hal.archives-ouvertes.fr/pasteur-01352619

D. Kearns, A field guide to bacterial swarming motility, Nature Reviews Microbiology, vol.33, issue.9, pp.634-644, 2010.
DOI : 10.1038/nrmicro2405

D. Maayer, P. Chan, W. Venter, S. Toth, I. Birch et al., Genome Sequence of Pantoea ananatis LMG20103, the Causative Agent of Eucalyptus Blight and Dieback, Journal of Bacteriology, vol.192, issue.11, pp.2936-293700060, 2010.
DOI : 10.1128/JB.00060-10

D. Maayer, P. Chan, W. Rezzonico, F. Bühlmann, A. Venter et al., Complete Genome Sequence of Clinical Isolate Pantoea ananatis LMG 5342, Journal of Bacteriology, vol.194, issue.6, pp.1615-161606715, 2012.
DOI : 10.1128/JB.06715-11

A. Sillo, J. Matthias, R. Konertz, S. Bozzaro, and L. Eichinger, Salmonella typhimurium is pathogenic for Dictyostelium cells and subverts the starvation response, Cellular Microbiology, vol.1469, issue.11, pp.1793-1811, 2011.
DOI : 10.1111/j.1462-5822.2011.01662.x

D. Smith, M. Kirzinger, and J. Stavrinides, Draft Genome Sequence of the Antibiotic-Producing Epiphytic Isolate Pantoea ananatis BRT175, Genome Announcements, vol.1, issue.6, pp.902-915, 2013.
DOI : 10.1128/genomeA.00902-13

S. Pukatzki, A. Ma, D. Sturtevant, B. Krastins, D. Sarracino et al., Identification of a conserved bacterial protein secretion system in Vibrio cholerae using the Dictyostelium host model system, Proceedings of the National Academy of Sciences, vol.103, issue.5, pp.1528-1533, 2006.
DOI : 10.1073/pnas.0510322103

E. Pradel, Y. Zhang, N. Pujol, T. Matsuyama, C. Bargmann et al., Detection and avoidance of a natural product from the pathogenic bacterium Serratia marcescens by Caenorhabditis elegans, Proceedings of the National Academy of Sciences, vol.104, issue.7, pp.2295-2300, 2007.
DOI : 10.1073/pnas.0610281104

URL : https://hal.archives-ouvertes.fr/hal-00165358

L. Antunes, R. Ferreira, M. Buckner, and B. Finlay, Quorum sensing in bacterial virulence, Microbiology, vol.156, issue.8, pp.2271-2282, 2010.
DOI : 10.1099/mic.0.038794-0

J. Pearson, E. Pesci, and B. Iglewski, Roles of Pseudomonas aeruginosa las and rhl quorum-sensing systems in control of elastase and rhamnolipid biosynthesis genes., Journal of Bacteriology, vol.179, issue.18, pp.5756-5767, 1997.
DOI : 10.1128/jb.179.18.5756-5767.1997

A. Deep, U. Chaudhary, and V. Gupta, Quorum sensing and bacterial pathogenicity: From molecules to disease, Journal of Laboratory Physicians, vol.3, issue.1, pp.4-11, 2011.
DOI : 10.4103/0974-2727.78553

L. Zulianello, C. Canard, T. Köhler, D. Caille, J. Lacroix et al., Rhamnolipids Are Virulence Factors That Promote Early Infiltration of Primary Human Airway Epithelia by Pseudomonas aeruginosa, Infection and Immunity, vol.74, issue.6, pp.3134-3147, 2006.
DOI : 10.1128/IAI.01772-05

P. Jensen, T. Bjarnsholt, R. Phipps, T. Rasmussen, H. Calum et al., Rapid necrotic killing of polymorphonuclear leukocytes is caused by quorum-sensing-controlled production of rhamnolipid by Pseudomonas aeruginosa, Microbiology, vol.153, issue.5, pp.1329-1338003863, 2006.
DOI : 10.1099/mic.0.2006/003863-0

S. Häussler, M. Rohde, N. Von-neuhoff, M. Nimtz, and I. Steinmetz, Structural and Functional Cellular Changes Induced by Burkholderia pseudomallei Rhamnolipid, Infection and Immunity, vol.71, issue.5, pp.2970-2975, 2003.
DOI : 10.1128/IAI.71.5.2970-2975.2003

U. Ochsner, A. Fiechter, and J. Reiser, Isolation, characterization, and expression in Escherichia coli of the Pseudomonas aeruginosa rhlAB genes encoding a rhamnosyltransferase involved in rhamnolipid biosurfactant synthesis, J Biol Chem, vol.269, pp.19787-19795, 1994.

D. Dubeau, E. Déziel, D. Woods, and F. Lépine, Burkholderia thailandensis harbors two identical rhl gene clusters responsible for the biosynthesis of rhamnolipids, BMC Microbiology, vol.9, issue.1, 2009.
DOI : 10.1186/1471-2180-9-263

R. Rahim, U. Ochsner, C. Olvera, M. Graninger, P. Messner et al., Cloning and functional characterization of the Pseudomonas aeruginosa rhlC gene that encodes rhamnosyltransferase 2, an enzyme responsible for di-rhamnolipid biosynthesis, Molecular Microbiology, vol.58, issue.3, pp.708-718, 2001.
DOI : 10.1046/j.1365-2958.2001.02420.x

E. Déziel, F. Lépine, S. Milot, and R. Villemur, Mass spectrometry monitoring of rhamnolipids from a growing culture of Pseudomonas aeruginosa strain 57RP, Biochimica et Biophysica Acta (BBA) - Molecular and Cell Biology of Lipids, vol.1485, issue.2-3, pp.145-152, 2000.
DOI : 10.1016/S1388-1981(00)00039-1

J. Tremblay, A. Richardson, F. Lépine, and E. Déziel, Self-produced P. ananatis Grazing Resistance, pp.75-90, 2007.

H. Brüssow, Bacteria between protists and phages: from antipredation strategies to the evolution of pathogenicity, Molecular Microbiology, vol.64, issue.3, pp.583-589, 2007.
DOI : 10.1111/j.1365-2958.2007.05826.x

C. Matz and S. Kjelleberg, Off the hook ??? how bacteria survive protozoan grazing, Trends in Microbiology, vol.13, issue.7, pp.302-307, 2005.
DOI : 10.1016/j.tim.2005.05.009

S. Brown, D. Cornforth, and N. Mideo, Evolution of virulence in opportunistic pathogens: generalism, plasticity, and control, Trends in Microbiology, vol.20, issue.7, pp.336-342, 2012.
DOI : 10.1016/j.tim.2012.04.005

S. Adiba, C. Nizak, M. Van-baalen, E. Denamur, and F. Depaulis, From Grazing Resistance to Pathogenesis: The Coincidental Evolution of Virulence Factors, PLoS ONE, vol.66, issue.8, 2010.
DOI : 10.1371/journal.pone.0011882.t002

URL : https://hal.archives-ouvertes.fr/hal-00528524

P. Fey, A. Kowal, P. Gaudet, K. Pilcher, and R. Chisholm, Protocols for growth and development of Dictyostelium discoideum, Nature Protocols, vol.21, issue.6, pp.1307-1316, 2007.
DOI : 10.1038/nprot.2007.178

M. Sussman, Chapter 14 Biochemical and Genetic Methods in the Study of Cellular Slime Mold Development, Methods in cell physiology, pp.397-410, 1966.
DOI : 10.1016/S0091-679X(08)62146-9

H. Ochman, A. Gerber, and D. Hartl, Genetic applications of an inverse polymerase chain reaction, Genetics, vol.120, pp.621-623, 1988.

L. Guerra-santos, O. Kappeli, and A. Fiechter, Pseudomonas aeruginosa biosurfactant production in continuous culture with glucose as carbon source, Appl Environ Microbiol, vol.48, pp.301-305, 1984.

A. Dinno and . Dunn, Test: Dunn's test of multiple comparisons using rank sums. R package version 1.2.4. R Foundation for Statistical Computing

A. Abdel-mawgoud, F. Lépine, and E. Déziel, Liquid Chromatography/Mass Spectrometry for the Identification and Quantification of Rhamnolipids, Methods Mol Biol, vol.1149, pp.359-373978, 2014.
DOI : 10.1007/978-1-4939-0473-0_30

URL : https://hal.archives-ouvertes.fr/pasteur-01134344

K. Tamura, G. Stecher, D. Peterson, A. Filipski, and S. Kumar, MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0, Molecular Biology and Evolution, vol.30, issue.12, pp.2725-2729, 2013.
DOI : 10.1093/molbev/mst197

B. Hall, H. Acar, A. Nandipati, and M. Barlow, Growth Rates Made Easy, Molecular Biology and Evolution, vol.31, issue.1, pp.232-238, 2014.
DOI : 10.1093/molbev/mst187

R. Team, R: a language and environment for statistical computing. R Foundation for Statistical Computing, 2014.

H. Wickham, ggplot2: elegant graphics for data analysis, 2009.

B. Kessler, D. Lorenzo, V. Timmis, and K. , A general system to integratelacZ fusions into the chromosomes of gram-negative eubacteria: regulation of thePm promoter of theTOL plasmid studied with all controlling elements in monocopy, Molecular and General Genetics MGG, vol.124, issue.1-2, pp.293-301, 1992.
DOI : 10.1007/BF00587591

P. Fey, R. Dodson, S. Basu, and R. Chisholm, One Stop Shop for Everything Dictyostelium: dictyBase and the Dicty Stock Center in 2012, Methods Mol Biol, vol.983, pp.59-92978, 2013.
DOI : 10.1007/978-1-62703-302-2_4