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Caracterizacion Molecular Y Funcional De Bacterias Del Genero Caracterización molecular y funcional de bacterias del género Azotobacter aisladas de suelos de la República Argentina. El rol de las auxinas en la respuesta a la inoculación de trigo. Tesis presentada para optar al título de Magister de la Universidad de Buenos Aires, Área Producción Vegetal con orientación en cultivos extensivos Esteban Julián Rubio Ingeniero Agrónomo-Universidad Nacional de Luján-2003 Lugar de trabajo: Instituto Nacional de Tecnología Agropecuaria (INTA) Escuela para Graduados Ing. Agr. Alberto Soriano Facultad de Agronomía – Universidad de Buenos Aires COMITÉ CONSEJERO Directora de tesis Olga Susana Correa Ingeniera Agrónoma Magister Scientiae Consejeros de Estudios Fabricio Darío Cassán Microbiólogo (Universidad Nacional de Río Cuarto) Doctor en Ciencias Biológicas (Universidad Nacional de Río Cuarto) Alejandro Perticari Ingeniero Agrónomo (Universidad de Morón) JURADO DE TESIS Directora de tesis Olga Susana Correa Ingeniera Agrónoma (Universidad de Buenos Aires) Magister Scientiae (Universidad de Buenos Aires) JURADO José Alfredo Curá Ingeniero Agrónomo (Universidad Nacional de Mar del Plata) Doctor en Ciencias (Área biología) (Universidad Nacional de Mar del Plata) JURADO Román Augusto Serrago Ingeniero Agrónomo (Universidad de Buenos Aires) Doctor en Ciencias Agropecuarias (Universidad de Buenos Aires) Fecha de defensa de la tesis: 2 de agosto de 2011 iii Dedicada a Gimena y a mi hijo Matías iv Agradecimientos A mi directora y mis consejeros de tesis, a las Dras Marcela Montecchia e Irma Roberts, a mis compañeros del laboratorio y a Gimena. v “Declaro que el material incluido en esta tesis es, a mi mejor saber y entender, original, producto de mi propio trabajo (salvo en la medida en que se identifique explícitamente las contribuciones de otros), y que este material no lo he presentado, en forma parcial o total, como una tesis en ésta u otra institución.” Esteban Julián Rubio vi ÍNDICE GENERAL Agradecimientos…………………………………………………………….…………iv Declaración……………………………………………………………………………..v Índice de Cuadros……………………..………………………….….…..…..….........viii Índice de Figuras………..…………………………………………...………..….........ix Resumen……………………………………………………..………………................xi Abstract…………………………………...……………………………….…..............xii 1. Introducción General……………………………..………...……………..…...........1 1.1. Objetivos generales…………………….…….…….………………..………….…5 Capítulo 1: “Diversidad de Azotobacter en suelos argentinos. Aislamiento y caracterización genotípica y fenotípica de los aislamientos”...………………...….....7 Introducción………………………………………………………………….......9 Objetivos e hipótesis……………..….………………………..….……………..11 Materiales y Métodos…………………………………….………………….….12 Resultados…………………………….…...…………………………………....17 Discusión……………………………….…..…………………………..............33 Capítulo 2: “Evaluación in vitro de características de cepas de Azotobacter relacionadas con la promoción del crecimiento vegetal”………………………...…35 Introducción……………..……………………………………….………….....37 Objetivos e hipótesis.……………………….…………...………….……….….40 Materiales y Métodos………...………….………………..……….…………...41 Resultados……………….………..…..………………………….…………….44 Discusión…………………..……………………..……………….…………....54 vii Capítulo 3: “Efecto de Azotobacter sobre la germinación y el crecimiento temprano de plantas de trigo”………………………...…………….……………………..……..57 Introducción………….……...…….…………………………………………....59 Objetivos e hipótesis.……………….………......………………………………60 Materiales y Métodos…………………………………..………………….…....61 Resultados…………………….……………………………………………..….63 Discusión……………………..………………...………………...………….....67 Capítulo 4: La producción de auxinas por Azotobacter como mecanismo responsable de la promoción del crecimiento temprano de trigo..............................71 Introducción…………..……….….…………………..………………………...73 Objetivos e hipótesis.………………..………………….………………..……..73 Materiales y Métodos……………………..…….………….………………..….74 Resultados…………….……………………………..……………………….....75 Discusión………………..…………………………...………………………....80 Discusión y conclusiones finales………………...………….…..…...……………..…83 Bibliografía……………..…………………..…………….……………………...…….87 Apéndice……………..………………………..……..…………………………..…….99 viii ÍNDICE DE CUADROS Cuadro 1. Muestras de suelo empleadas para el aislamiento de Azotobacter…………..19 Cuadro 2. Morfología de las colonias de los aislamientos de Azotobacter crecidas en medio LG luego de 7 días………………………………………………………………27 Cuadro 3. Velocidad específica de crecimiento (µ) tiempo de generación (g), ufc.ml-1 a las 11 y 185 h de crecimiento y producción de auxinas por parte de cepas de Azotobacter………………………………………………………....……………………………51 Cuadro 4. Producción de ácido-3-indolacético (AIA), ácido giberélico (GA3) y zeatina (Z), luego de 5 días (cepas AT18 y AT42) u 8 días de crecimiento (cepas AT19, AT25, AT31 y AT37)…...………………………………..…………………………………….52 ix ÍNDICE DE FIGURAS Figura 1. Dendrograma del análisis de los perfiles de rep-PCR de los aislamientos….23 Figura 2. Dendrograma del ARDRA con la enzima HhaI de las cepas eleccionadas………………………………………………………………………..…...25 Figura 3. Colonias de aislamientos Azotobacter desarrolladas en medio LG luego de 7 días de crecimiento……………………………………………………………….…….29 Figura 4. Tinción de Gram observada con microscopio óptico de cultivos puros de cepas Azotobacter luego de 48 hs de crecimiento en medio LG sólido……………….30 Figura 5. Producción de pigmentos no difusibles de cepas de Azotobacter en medio de Norris y Jensen luego de 10 días.....................................................................................31 Figura 6. Tinciones de Gram observadas con microscopio óptico de los quistes producidos por cepas de Azotobacter luego de 7 días de incubación en medio Burk con 0,3% de butanol como fuente de carbono……………………………………...…….…32 Figura 7 a. Logaritmo del número de unidades formadoras de colonia por ml de cultivo (ufc.ml-1) y producción de auxinas de cepas de Azotobacter crecidas en cultivos líquidos, en función del tiempo………………………………..........................45 Figura 7 b. Logaritmo del número de unidades formadoras de colonia por ml de cultivo (ufc.ml-1) y producción auxinas de cepas de Azotobacter crecidas en cultivos líquidos, en función del tiempo….…………….…………...…………........46 Figura 7 c. Logaritmo del número de unidades formadoras de colonia por ml de cultivo (ufc.ml-1) y producción auxinas de cepas de Azotobacter crecidas en cultivos líquidos, en función del tiempo……………………...……….….................47 Figura 8. Ensayos de producción de sideróforos……………….………...……...…….49 Figura 9. Ensayos de solubilización de fosfatos………….…………………….……...50 Figura 10. Comparación entre la producción de AIA determinada por LC-MS y la producción de auxinas medida por colorimetría (Salkowsky) para los mismos cultivos………………………………..………………………………..……..52 Figura 11. Actividad de nitrogenasa por la técnica de reducción de acetileno de cepas de Azotobacter…………………………..……..……………………………………..53 Figura 12. Velocidad germinativa de plántulas de trigo inoculadas con cepas de Azotobacter de baja (AT18 y AT37, AT42) y de alta producción de auxinas x (AT19, AT25 y AT31)……………...…….………………………………………….....63 Figura 13. Número de raíces seminales de plántulas de trigo a los 8 días de la inoculación con cepas de Azotobacter de baja (AT18 y AT37, AT42) y de alta producción de auxinas (AT19, AT25 y AT31)…….…......……………..…….……..….64 Figura 14. Peso seco de la raíz (a) y del tallo (b) de plántulas de trigo a los 8 días de la inoculación con cepas de Azotobacter de baja (AT18 y AT37, AT42) y de alta producción de auxinas (AT19, AT25 y AT31)…………………………….……....64 Figura 15. Unidades formadoras de colonia por raíz de plántulas de trigo a los 8 días de la inoculación con cepas de Azotobacter……………..............…………………….65 Figura 16. Longitud del la raíz (cm) de plántulas de trigo a los 8 días de la aplicación de cultivos bacterianos completos (C) y sobrenadantes (S) de cepas de Azotobacter de alta (AT19 y AT25), de baja producción de auxinas (AT18) y de soluciones de AIA de baja (AIAb), media (AIAm) y alta (AIAa) concentración de AI…………………...…..75 Figura 17. Longitud (I) y peso seco del tallo (II) de plantas de trigo a los 8 días de la aplicación de cultivos bacterianos completos (C) o sobrenadantes libres de células (S) del cultivo de cepas de Azotobacter de alta (AT19 y AT25) y baja producción de AIA (AT18) y de soluciones AIA de concentraciones baja (AIAb), media (AIAm) y alta (AIAa)…………………………………………………………………………..............76 Figura 18. Número de raíces seminales de plántulas de trigo a los 8 días de la aplicación de cultivos bacterianos completos (C) o sobrenadantes libres de células (S) del cultivo de cepas de Azotobacter de alta (AT19 y AT25) y baja producción de AIA (AT18) y de soluciones AIA de concentraciones baja (AIAb), media (AIAm) y alta (AIAa)………………………………………………………...……………….……..…77 Figura 19. Plántulas de trigo a las 24 h del tratamiento con cultivos completos o sobrenadantes de las cepas AT18, AT19, y AT25 o con soluciones de 2 µg.ml-1 (AIAb), 14 µg.ml-1 (AIAm) y 26 µg.ml-1 (AIAa)..........…………….…...78 Figura 20. Plántulas de trigo de 4 días tratadas con soluciones de 2 µ.ml-1 (AIAb), 14 µ.ml-1 (AIAb) y 26 µ.ml-1 (AIAa) y con sobrenadantes del cultivo de las cepas AT18 y AT19………………………………..…………………………………...…………...…78 Figura 21. Raíces de plántulas de trigo de 4 días tratadas con soluciones de 2 µ.ml-1 (AIAb), 14 µ.ml-1 (AIAb) y 26 µ.ml-1 (AIAa) y con sobrenadantes del cultivo de las cepas AT18 y AT19………………………………...……..…………….……....79 xi Caracterización molecular y funcional de bacterias del género Azotobacter aisladas de suelos de la República Argentina. El
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