TGCAAGTCGAACGAAGTCTTCGGACTTAGTGGCGCACGGGTGCGTAACGCGTGGGAATCTGCCCTTGGGTACGGAATAACTCAGAGAAATTTGT GCTAATACCGTATAATGTCTTCGGACCAAAGATTTATCGCCCAAGGATGAGCCCGCGTAAGATTAGCTAGTTGGTGGGGTAAAAGCCTACCAAG GCGACGATCTTTAGCTGGTCTGAGAGGATGATCAGCCACACTGGGACTGAGACACGGCCCAGACTCCTACGGGAGGCAGCAGTGGGGAATATT GGACAATGGGCGAAAGCCTGATCCAGCAATGCCGCGTGAGTGATGAAGGCCTTAGGGTTGTAAAGCTCTTTTACCCGGGATGATAATGACAGTA CCGGGAGAATAAGCTCCGGCTAACTCCGTGCCAGCAGCCGCGGTAATACGGAGGGAGCTAGCGTTGTTCGGAATTACTGGGCGTAAAGCGCGC GTAGGCGGTTTTTTAAGTCAGAGGTGAAAGCCCAGTGCTCAACACTGGAACTGCCTTTGAAACTGGAAAACTTGAATCTTGGAGAGGTCAGTGG AATTCCGAGTGTAGAGGTGAAATTCGTAGATATTCGGAAGAACACCAGTGGCGAAGGCGACTGACTGGACAAGTATTGACGCTGAGGTGCGAA AGCGTGGGGAGCAAACAGGATTAGATACCCTGGTAGTCCACGCCGTAAACGATGATAACTAGCTGTCCGGGTTCATAGAACTTGGGTGGCGCAGDoctoral Dissertation CTAACGCATTAAGTTATCCGCCTGGGGAGTACGGTCGCAAGATTAAAACTCAAAGGAATTGACGGGGGCCTGCACAAGCGGTGGAGCATGTGG TTTAATTCGAAGCAACGCGCAGAACCTTACCAGCGTTTGACATCCTGATCGCGGATTAGAGAGATCTTTTCCTTCAGTTCGGCTGGATCAGTGACA GGTGCTGCATGGCTGTCGTCAGCTCGTGTCGTGAGATGTTGGGTTAAGTCCCGCAACGAGCGCAACCCTCATCCCTAGTTGCCATCATTAAGTTG GGCACTCTAAGGAAACTGCCGGTGATAAGCCGGAGGAAGGTGGGGATGACGTCAAGTCCTCATGGCCCTTACGCGCTGGGCTACACACGTGCT ACAATGGCAACTACAGTGGGCAGCAACCTCGCGAGGGGTAGCTAATCTCCAAAAGTTGTCTCAGTTCGGATTGTTCTCTGCAACTCGAGAGCATG AAGGCGGAATCGCTAGTAATCGCGGATCAGCATGCCGCGGTGAATACGTTCCCAGGCCTTGTACACACCGCCCGTCACACCATGGGAGTTGGTTBiodegradation of microcystins by TCACCCGAAGGCAGTGCTCTAACCCGCAATGCAAGTCGAACGAAGTCTTCGGACTTAGTGGCGCACGGGTGCGTAACGCGTGGGAATCTGCCCT TGGGTACGGAATAACTCAGAGAAATTTGTGCTAATACCGTATAATGTCTTCGGACCAAAGATTTATCGCCCAAGGATGAGCCCGCGTAAGATTAG CTAGTTGGTGGGGTAAAAGCCTACCAAGGCGACGATCTTTAGCTGGTCTGAGAGGATGATCAGCCACACTGGGACTGAGACACGGCCCAGACTC CTACGGGAGGCAGCAGTGGGGAATATTGGACAATGGGCGAAAGCCTGATCCAGCAATGCCGCGTGAGTGATGAAGGCCTTAGGGTTGTAAAGCfreshwater bacteria: TCTTTTACCCGGGATGATAATGACAGTACCGGGAGAATAAGCTCCGGCTAACTCCGTGCCAGCAGCCGCGGTAATACGGAGGGAGCTAGCGTTG TTCGGAATTACTGGGCGTAAAGCGCGCGTAGGCGGTTTTTTAAGTCAGAGGTGAAAGCCCAGTGCTCAACACTGGAACTGCCTTTGAAACTGGA AAACTTGAATCTTGGAGAGGTCAGTGGAATTCCGAGTGTAGAGGTGAAATTCGTAGATATTCGGAAGAACACCAGTGGCGAAGGCGACTGACT GGACAAGTATTGACGCTGAGGTGCGAAAGCGTGGGGAGCAAACAGGATTAGATACCCTGGTAGTCCACGCCGTAAACGATGATAACTAGCTGT CCGGGTTCATAGAACTTGGGTGGCGCAGCTAACGCATTAAGTTATCCGCCTGGGGAGTACGGTCGCAAGATTAAAACTCAAAGGAATTGACGGGNew genetic and ecological perspectives GGCCTGCACAAGCGGTGGAGCATGTGGTTTAATTCGAAGCAACGCGCAGAACCTTACCAGCGTTTGACATCCTGATCGCGGATTAGAGAGATCT TTTCCTTCAGTTCGGCTGGATCAGTGACAGGTGCTGCATGGCTGTCGTCAGCTCGTGTCGTGAGATGTTGGGTTAAGTCCCGCAACGAGCGCAAC CCTCATCCCTAGTTGCCATCATTAAGTTGGGCACTCTAAGGAAACTGCCGGTGATAAGCCGGAGGAAGGTGGGGATGACGTCAAGTCCTCATGG CCCTTACGCGCTGGGCTACACACGTGCTACAATGGCAACTACAGTGGGCAGCAACCTCGCGAGGGGTAGCTAATCTCCAAAAGTTGTCTCAGTTC GGATTGTTCTCTGCAACTCGAGAGCATGAAGGCGGAATCGCTAGTAATCGCGGATCAGCATGCCGCGGTGAATACGTTCCCAGGCCTTGTACAC ACCGCCCGTCACACCATGGGAGTTGGTTTCACCCGAAGGCAGTGCTCTAACCCGCAATGCAAGTCGAACGAAGTCTTCGGACTTAGTGGCGCAC GGGTGCGTAACGCGTGGGAATCTGCCCTTGGGTACGGAATAACTCAGAGAAATTTGTGCTAATACCGTATAATGTCTTCGGACCAAAGATTTATC GCCCAAGGATGAGCCCGCGTAAGATTAGCTAGTTGGTGGGGTAAAAGCCTACCAAGGCGACGATCTTTAGCTGGTCTGAGAGGATGATCAGCCA CACTGGGACTGAGACACGGCCCAGACTCCTACGGGAGGCAGCAGTGGGGAATATTGGACAATGGGCGAAAGCCTGATCCAGCAATGCCGCGTG AGTGATGAAGGCCTTAGGGTTGTAAAGCTCTTTTACCCGGGATGATAATGACAGTACCGGGAGAATAAGCTCCGGCTAACTCCGTGCCAGCAGC CGCGGTAATACGGAGGGAGCTAGCGTTGTTCGGAATTACTGGGCGTAAAGCGCGCGTAGGCGGTTTTTTAAGTCAGAGGTGAAAGCCCAGTGCT CAACACTGGAACTGCCTTTGAAACTGGAAAACTTGAATCTTGGAGAGGTCAGTGGAATTCCGAGTGTAGAGGTGAAATTCGTAGATATTCGGAA GAACACCAGTGGCGAAGGCGACTGACTGGACAAGTATTGACGCTGAGGTGCGAAAGCGTGGGGAGCAAACAGGATTAGATACCCTGGTAGTCC ACGCCGTAAACGATGATAACTAGCTGTCCGGGTTCATAGAACTTGGGTGGCGCAGCTAACGCATTAAGTTATCCGCCTGGGGAGTACGGTCGCA AGATTAAAACTCAAAGGAATTGACGGGGGCCTGCACAAGCGGTGGAGCATGTGGTTTAATTCGAAGCAACGCGCAGAACCTTACCAGCGTTTGA CATCCTGATCGCGGATTAGAGAGATCTTTTCCTTCAGTTCGGCTGGATCAGTGACAGGTGCTGCATGGCTGTCGTCAGCTCGTGTCGTGAGATGTT GGGTTAAGTCCCGCAACGAGCGCAACCCTCATCCCTAGTTGCCATCATTAAGTTGGGCACTCTAAGGAAACTGCCGGTGATAAGCCGGAGGAAG GTGGGGATGACGTCAAGTCCTCATGGCCCTTACGCGCTGGGCTACACACGTGCTACAATGGCAACTACAGTGGGCAGCAACCTCGCGAGGGGTA GCTAATCTCCAAAAGTTGTCTCAGTTCGGATTGTTCTCTGCAACTCGAGAGCATGAAGGCGGAATCGCTAGTAATCGCGGATCAGCATGCCGCGG TGAATACGTTCCCAGGCCTTGTACACACCGCCCGTCACACCATGGGAGTTGGTTTCACCCGAAGGCAGTGCTCTAACCCGCAATGCAAGTCGAAC GAAGTCTTCGGACTTAGTGGCGCACGGGTGCGTAACGCGTGGGAATCTGCCCTTGGGTACGGAATAACTCAGAGAAATTTGTGCTAATACCGTA TAATGTCTTCGGACCAAAGATTTATCGCCCAAGGATGAGCCCGCGTAAGATTAGCTAGTTGGTGGGGTAAAAGCCTACCAAGGCGACGATCTTTA GCTGGTCTGAGAGGATGATCAGCCACACTGGGACTGAGACACGGCCCAGACTCCTACGGGAGGCAGCAGTGGGGAATATTGGACAATGGGCGA AAGCCTGATCCAGCAATGCCGCGTGAGTGATGAAGGCCTTAGGGTTGTAAAGCTCTTTTACCCGGGATGATAATGACAGTACCGGGAGAATAAG CTCCGGCTAACTCCGTGCCAGCAGCCGCGGTAATACGGAGGGAGCTAGCGTTGTTCGGAATTACTGGGCGTAAAGCGCGCGTAGGCGGTTTTTT AAGTCAGAGGTGAAAGCCCAGTGCTCAACACTGGAACTGCCTTTGAAACTGGAAAACTTGAATCTTGGAGAGGTCAGTGGAATTCCGAGTGTAG AGGTGAAATTCGTAGATATTCGGAAGAACACCAGTGGCGAAGGCGACTGACTGGACAAGTATTGACGCTGAGGTGCGAAAGCGTGGGGAGCA AACAGGATTAGATACCCTGGTAGTCCACGCCGTAAACGATGATAACTAGCTGTCCGGGTTCATAGAACTTGGGTGGCGCAGCTAACGCATTAAG TTATCCGCCTGGGGAGTACGGTCGCAAGATTAAAACTCAAAGGAATTGACGGGGGCCTGCACAAGCGGTGGAGCATGTGGTTTAATTCGAAGCA ACGCGCAGAACCTTACCAGCGTTTGACATCCTGATCGCGGATTAGAGAGATCTTTTCCTTCAGTTCGGCTGGATCAGTGACAGGTGCTGCATGGC TGTCGTCAGCTCGTGTCGTGAGATGTTGGGTTAAGTCCCGCAACGAGCGCAACCCTCATCCCTAGTTGCCATCATTAAGTTGGGCACTCTAAGGA AACTGCCGGTGATAAGCCGGAGGAAGGTGGGGATGACGTCAAGTCCTCATGGCCCTTACGCGCTGGGCTACACACGTGCTACAATGGCAACTAC AGTGGGCAGCAACCTCGCGAGGGGTAGCTAATCTCCAAAAGTTGTCTCAGTTCGGATTGTTCTCTGCAACTCGAGAGCATGAAGGCGGAATCGC TAGTAATCGCGGATCAGCATGCCGCGGTGAATACGTTCCCAGGCCTTGTACACACCGCCCGTCACACCATGGGAGTTGGTTTCACCCGAAGGCAG TGCTCTAACCCGCAATGCAAGTCGAACGAAGTCTTCGGACTTAGTGGCGCACGGGTGCGTAACGCGTGGGAATCTGCCCTTGGGTACGGAATAA CTCAGAGAAATTTGTGCTAATACCGTATAATGTCTTCGGACCAAAGATTTATCGCCCAAGGATGAGCCCGCGTAAGATTAGCTAGTTGGTGGGGT AAAAGCCTACCAAGGCGACGATCTTTAGCTGGTCTGAGAGGATGATCAGCCACACTGGGACTGAGACACGGCCCAGACTCCTACGGGAGGCAG CAGTGGGGAATATTGGACAATGGGCGAAAGCCTGATCCAGCAATGCCGCGTGAGTGATGAAGGCCTTAGGGTTGTAAAGCTCTTTTACCCGGGA TGATAATGACAGTACCGGGAGAATAAGCTCCGGCTAACTCCGTGCCAGCAGCCGCGGTAATACGGAGGGAGCTAGCGTTGTTCGGAATTACTGG GCGTAAAGCGCGCGTAGGCGGTTTTTTAAGTCAGAGGTGAAAGCCCAGTGCTCAACACTGGAACTGCCTTTGAAACTGGAAAACTTGAATCTTG GAGAGGTCAGTGGAATTCCGAGTGTAGAGGTGAAATTCGTAGATATTCGGAAGAACACCAGTGGCGAAGGCGACTGACTGGACAAGTATTGAC GCTGAGGTGCGAAAGCGTGGGGAGCAAACAGGATTAGATACCCTGGTAGTCCACGCCGTAAACGATGATAACTAGCTGTCCGGGTTCATAGAA CTTGGGTGGCGCAGCTAACGCATTAAGTTATCCGCCTGGGGAGTACGGTCGCAAGATTAAAACTCAAAGGAATTGACGGGGGCCTGCACAAGC GGTGGAGCATGTGGTTTAATTCGAAGCAACGCGCAGAACCTTACCAGCGTTTGACATCCTGATCGCGGATTAGAGAGATCTTTTCCTTCAGTTCG GCTGGATCAGTGACAGGTGCTGCATGGCTGTCGTCAGCTCGTGTCGTGAGATGTTGGGTTAAGTCCCGCAACGAGCGCAACCCTCATCCCTAGTT GCCATCATTAAGTTGGGCACTCTAAGGAAACTGCCGGTGATAAGCCGGAGGAAGGTGGGGATGACGTCAAGTCCTCATGGCCCTTACGCGCTGG GCTACACACGTGCTACAATGGCAACTACAGTGGGCAGCAACCTCGCGAGGGGTAGCTAATCTCCAAAAGTTGTCTCAGTTCGGATTGTTCTCTGC AACTCGAGAGCATGAAGGCGGAATCGCTAGTAATCGCGGATCAGCATGCCGCGGTGAATACGTTCCCAGGCCTTGTACACACCGCCCGTCACAC CATGGGAGTTGGTTTCACCCGAAGGCAGTGCTCTAACCCGCAATGCAAGTCGAACGAAGTCTTCGGACTTAGTGGCGCACGGGTGCGTAACGCG TGGGAATCTGCCCTTGGGTACGGAATAACTCAGAGAAATTTGTGCTAATACCGTATAATGTCTTCGGACCAAAGATTTATCGCCCAAGGATGAGC CCGCGTAAGATTAGCTAGTTGGTGGGGTAAAAGCCTACCAAGGCGACGATCTTTAGCTGGTCTGAGAGGATGATCAGCCACACTGGGACTGAGA CACGGCCCAGACTCCTACGGGAGGCAGCAGTGGGGAATATTGGACAATGGGCGAAAGCCTGATCCAGCAATGCCGCGTGAGTGATGAAGGCCTMaría Ángeles Lezcano Vega TAGGGTTGTAAAGCTCTTTTACCCGGGATGATAATGACAGTACCGGGAGAATAAGCTCCGGCTAACTCCGTGCCAGCAGCCGCGGTAATACGGA GGGAGCTAGCGTTGTTCGGAATTACTGGGCGTAAAGCGCGCGTAGGCGGTTTTTTAAGTCAGAGGTGAAAGCCCAGTGCTCAACACTGGAACTG CCTTTGAAACTGGAAAACTTGAATCTTGGAGAGGTCAGTGGAATTCCGAGTGTAGAGGTGAAATTCGTAGATATTCGGAAGAACACCAGTGGCGMadrid, 2017 AAGGCGACTGACTGGACAAGTATTGACGCTGAGGTGCGAAAGCGTGGGGAGCAAACAGGATTAGATACCCTGGTAGTCCACGCCGTAAACGAT GATAACTAGCTGTCCGGGTTCATAGAACTTGGGTGGCGCAGCTAACGCATTAAGTTATCCGCCTGGGGAGTACGGTCGCAAGATTAAAACTCAA AGGAATTGACGGGGGCCTGCACAAGCGGTGGAGCATGTGGTTTAATTCGAAGCAACGCGCAGAACCTTACCAGCGTTTGACATCCTGATCGCGG ATTAGAGAGATCTTTTCCTTCAGTTCGGCTGGATCAGTGACAGGTGCTGCATGGCTGTCGTCAGCTCGTGTCGTGAGATGTTGGGTTAAGTCCCG CAACGAGCGCAACCCTCATCCCTAGTTGCCATCATTAAGTTGGGCACTCTAAGGAAACTGCCGGTGATAAGCCGGAGGAAGGTGGGGATGACGT CAAGTCCTCATGGCCCTTACGCGCTGGGCTACACACGTGCTACAATGGCAACTACAGTGGGCAGCAACCTCGCGAGGGGTAGCTAATCTCCAAA Universidad Autónoma de Madrid Facultad de Ciencias Departamento de Biología Biodegradation of microcystins by freshwater bacteria: new genetic and ecological perspectives Biodegradación de microcistinas por bacterias de aguas dulces: nuevas perspectivas genéticas y ecológicas Doctoral Dissertation Tesis Doctoral MARÍA ÁNGELES LEZCANO VEGA Madrid, Junio 2017 Memoria presentada para optar al grado de Doctor en Microbiología María Ángeles Lezcano Vega Licenciada en Ciencias Ambientales Directores: Dra. Rehab El-Shehawy Investigadora Instituto IMDEA Agua Dr. Antonio Quesada del Corral Profesor titular Departamento de Biología Universidad Autónoma de Madrid Madrid a 26 de Junio de 2017 A mi familia Table of contents / Índice Acknowledgements / Agradecimientos ......................................................................... i Abstract / Resumen ..................................................................................................... v Abbreviations / Abreviaturas ..................................................................................... ix Figure index / Índice de figuras .................................................................................. xi Table index / Índice de tablas................................................................................... xiii Chapter 1 General introduction ................................................................................................... 3 1.1. Cyanobacteria