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Caracterización De Especies Fitopatógenas De CARACTERIZACIÓN DE ESPECIES FITOPATÓGENAS DE PYTHIUM Y PHYTOPHTHORA (PERONOSPOROMYCETES) EN CULTIVOS ORNAMENTALES DEL CINTURÓN VERDE LA PLATA-BUENOS AIRES Y OTRAS ÁREAS Y CULTIVOS DE INTERÉS TESIS PARA OPTAR AL TÍTULO DE DOCTOR EN CIENCIAS NATURALES FACULTAD DE CIENCIAS NATURALES Y MUSEO UNIVERSIDAD NACIONAL DE LA PLATA HEMILSE ELENA PALMUCCI DIRECTOR: ING. AGR. SILVIA WOLCAN CODIRECTOR: DRA MÓNICA STECIOW AÑO 2015 1 AGRADECIMIENTOS A la Facultad de Ciencias Naturales y Museo (FCNYM) por brindarme la posibilidad de realizar este trabajo A la Ing Agr Silvia Wolcan y a la Dra Mónica Steciow por sus sugerencias y comentarios en la ejecución y escritura de esta tesis. A la Dra Gloria Abad, investigadora líder en Oomycetes en el “USDA- Molecular Diagnostic Laboratory (MDL)”, por su invalorable y generosa colaboración en mi formación a través de sus conocimientos, por brindarme la posibilidad de llevar a cabo los trabajos moleculares en el MDL-Maryland-USA y apoyar mi participación en workshops internacionales y reuniones de la especialidad. A la Ing Wolcan por aportar su valiosa colección para realizar parte de las tareas de identificación y por los significativos aportes realizados desde su experiencia. A la Dra Gloria Abad, a la Ing. Agr. Silvia Wolcan, al Dr R. Dehley y al Ing. Agr. Carlos Carloni por la provisión de valiosas referencias bibliográficas. Al Ing Agr M Sc Pablo Grijalba por acompañarme en la ardua tarea de introducirnos al mundo de los Oomycetes a través de nuestras tesis doctorales y los proyectos de investigación que compartimos, y por sus consejos y colaboración en aspectos biomoleculares. A mis compañeros del Proyecto Ubacyt, Lic. Elisa Fantino, Cecilia Herrera y estudiantes tesistas quienes colaboraron en el mantenimiento de la colección, la ejecución de algunas pruebas de patogenicidad y apoyo técnico en tareas de laboratorio. A los productores que facilitaron mi ingreso a sus establecimientos o me enviaron muestras enfermas, a los Ings Agrs Daniel Morisigue y Oscar Herrera, a los viveros “Plugsystem” y “La Facultad”, por facilitarme plantas para la realización de las pruebas de patogenicidad. A la Dra Mary Palm, Directora del que fuera el “USDA- MDL-Maryland-USA”, por facilitar mi estadía de entrenamiento. Al Dr Seoghan Kant de la “Penn State University” por los insumos donados para parte de los estudios moleculares y al Dr Balci de la “University of Maryland” por facilitar instrumental óptico para toma de fotografías microscópicas en 2012. Muy especialmente a mi esposo y mi hija que me acompañaron estos años con su amor y paciencia. 2 INDICE GENERAL AGRADECIMIENTOS 2 RESUMEN 8 ABSTRACT 12 INTRODUCCIÓN Evolución del conocimiento de los géneros Pythium y Phytophthora 17 en Argentina 1. Hábito nutricional y sintomatología. 17 2. Ubicación taxonómica. 20 3. Caracterización morfológica. 24 4. Supervivencia y dispersión del inóculo. Relación con el manejo de la 27 enfermedad 5. Criterios de clasificación: morfológico, molecular, filogenético. 31 6. Antecedentes sobre trabajos de relevamiento. 36 7. Estudios efectuados en Argentina 37 HIPÓTESIS 40 OBJETIVOS GENERALES Y ESPECÍFICOS 40 CARACTERIZACIÓN DEL SECTOR FLORÍCOLA Y DE LA REGIÓN EN ESTUDIO. 1. Caracterización general 43 2. Condiciones ambientales: clima y suelo 46 MATERIALES Y MÉTODOS 1. Antecedentes sobre especies fitopatógenas de los géneros Pythium 49 y Phytophthora en Argentina. 2. Prospección de enfermedades en el Cinturón verde La Plata- 50 Buenos Aires y otras áreas de interés. 3. Técnicas de aislamiento, purificación y mantenimiento de la colección 53 3.1 Aislamientos a partir de material sintomático. 53 3.2. Aislamiento desde agua de riego. 54 3.3. Medios de cultivo. 55 3.4. Purificación de los aislamientos. 55 3.5. Mantenimiento de la colección. 56 3.6. Depósito de los aislamientos en Colecciones micológicas 56 y de secuencias en Bancos Genéticos 4. Identificación y descripción de los aislamientos por métodos 57 micológicos clásicos. 3 4.1. Caracterización de la colonia 57 4.2. Caracterización morfológica 57 4.2.1. Pythium spp. 57 4.2.2. Phytophthora spp. 58 59 4.2.3. Estimulación de estructuras reproductivas 4.2.4. Observación de estructuras con microscopio óptico 60 4.3. Determinación de temperaturas de crecimiento 61 4.4. Identificación taxonómica 61 5. Pruebas de patogenicidad 62 5.1. Depósito de bloques de agar con inóculo en heridas 62 5.2. Colocación de agar con inóculo en contacto con las raíces 63 5.3. Aporte de inóculo utilizando un soporte orgánico 64 5,4. Inoculación con agua de riego 65 5.5. Inoculaciones en otros hospedantes 65 6. Identificación y caracterización de los aislamientos por métodos 65 moleculares 6.1. Extracción del ADN 65 6.2. Amplificación del rADN 65 6.3. Purificación y secuenciación del rADN 66 6.4. Construcción de árboles filogenéticos 67 7. Aspectos referidos a la bibliografía utilizada en el presente estudio 67 RESULTADOS PARTE I. ANTECEDENTES SOBRE ESPECIES FITOPATÓGENAS DE OOMYCETES: STATUS DE LOS GÉNEROS PYTHIUM Y PHYTOPHTHORA EN ARGENTINA 1. Introducción 2. El género Pythium en Argentina 72 3. El género Phytophthora en Argentina 94 PARTE II. CARACTERIZACIÓN DE ESPECIES FITOPATÓGENAS DE PYTHIUM Y PHYTOPHTHORA INTRODUCCIÓN 119 GÉNERO PYTHIUM 122 4 123 ESPECIES CON ESPORANGIOS LOBULADOS O FILAMENTOSOS P. aphanidermatum 123 P. dissotocum 140 P. graminicola 150 ESPECIES CON ESPORANGIOS ESFÉRICOS, SUBESFÉRICOS O ELIPSOIDALES A. OOGONIOS LISOS P. cylindrosporum 160 P. sylvaticum 169 P. ultimum var. ultimum 179 P. ultimum var. sporangiiferum 187 B. OOGONIOS ORNAMENTADOS P. irregulare 192 P. spinosum 201 C .ÓRGANOS SEXUALES AUSENTES P. intermedium 208 P. splendens 217 NUEVAS ESPECIES DE PYTHIUM Pythium sp. nov. 1 224 Pythium sp. nov. 2 233 GÉNERO PHYTOPYTHIUM Phytopythium chamaehyphon 242 ESPECIES DE PHYTOPHTHORA IDENTIFICADAS ESPECIES HETEROTÁLICAS ESPORANGIOS PAPILADOS Ph. capsici 253 Ph. nicotianae= parasitica 267 ESPECIES HETEROTÁLICAS ESPORANGIOS NO PAPILADOS Ph. cinnamomi 293 Ph. cryptogea 310 Ph. taxon kelmania 322 FILOGENIA DE ESPECIES DE PHYTOPHTHORA PRESENTES 329 EN ARGENTINA TABLAS RESUMEN DE RESULTADOS OBTENIDOS 333 DISCUSIÓN GENERAL 344 CONCLUSIONES 371 BIBLIOGRAFÍA CITADA 378 ANEXO I (Antecedentes de Pythium spp. en Argentina) 424 ANEXO II (Antecedentes de Phytophthora spp. en Argentina) 431 5 ÍNDICE DE TABLAS 1: Especies patógenas de Pythium citadas en Argentina. relación hospedante- 73 patógeno, síntomas y localidades donde fueron halladas 2: Distribución de Pythium spp. en regiones geográficas de Argentina en los 87 años 2009 y 2014 3: Número de hospedantes afectados por especies de Pythium dentro 89 de cada tipo de cultivo en los recuentos de 2009 y de 2014 4: Especies de Pythium citadas para el Cinturón verde La Plata-Buenos Aires 93 hasta el año 2009. 5: Especies de Pythium identificadas durante el presente estudio (2009-2014). 93 6: Especies patógenas de Phytophthora citadas en Argentina: relación 95 hospedante-patógeno, síntomas y localización geográfica 7: Especies de Phytophthora aisladas de suelo o agua 108 8: Especies de Phytophthora citadas para el Cinturón verde La Plata-Buenos 108 Aires hasta el año 2009. 9: Especies de Phytophthora identificadas durante el presente estudio (2009- 109 2013). 10: Número de hospedantes afectados por especies de Phytophthora dentro de 111 cada tipo de cultivo en los recuentos de 2009 y de 2014. 11: Distribución de Phytophthora spp. en regiones geográficas de Argentina 113 (año 2014). 12: Crecimiento promedio de los aislamientos de Ph. nicotianae a 20 y 28 °C. 282 13: Características culturales, morfológicas y biométricas de los aislamientos 282 de Ph. nicotianae. 6 ÍNDICE DE FIGURAS Fig. 1: Cinturón verde La Plata-Buenos Aires 47 Fig. 2: Número de especies de Pythium/ provincia en los años 2009-2014. 88 Fig. 3: Número total de hospedantes/tipo de cultivo afectados por especies 90 de Pythium (evaluaciones en 2009 y en 2014). Fig. 4: Frecuencia relativa de especies de Pythium (2014). 91 Fig. 5: Frecuencia relativa de Pythium spp. que afectan cultivos ornamentales 92 (2014). Fig.6: Número de especies de Phytophthora/tipo de cultivo (año 2014). 112 Fig. 7: Número de de especies de Phytophthora/provincia (2014). 115 Fig. 8: Frecuencia relativa de especies de Phytophthora (2014) 116 Fig. 9: Frecuencia relativa de Phytophthora spp. que afectan cultivos 117 ornamentales (2014) Figs. 10-17: Pythium aphanidermatum 126 Figs.18-21: P. dissotocum 141 Figs. 22-26: P. graminicola 151 Figs. 27- 31: P. cylindrosporum 164 Figs. 32-36: P. sylvaticum 173 Figs. 37-39: P. ultimum var. ultimum 183 Fig. 40-41: P. ultimum var. sporangiiferum 190 Figs. 42-44: P. irregulare 198 Figs. 45-47: P. spinosum 202 Figs. 48- 51: P. intermedium 213 Figs. 52-53: P. splendens 222 Figs. 54-59: Pythium sp. nov.1 226 Figs. 60-65: Pythium sp. nov.2 235 Figs. 66-70: Phytopythium chamaehyphon 244 Figs. 71-76: Ph. capsici 261 Figs. 77-89 : Ph. nicotianae= parasitica 269 Figs. 90-95: Ph. cinnamomi 304 Figs. 96-98: Ph. cryptogea 313 Figs. 99- Ph. taxon kelmania 326 7 CARACTERIZACIÓN DE ESPECIES FITOPATÓGENAS DE PYTHIUM Y PHYTOPHTHORA (PERONOSPOROMYCETES) EN CULTIVOS ORNAMENTALES DEL CINTURÓN VERDE LA PLATA-BUENOS AIRES Y OTRAS ÁREAS Y CULTIVOS DE INTERÉS RESUMEN La diversidad de especies de Pythium y de Phytophthora conocidas en el mundo y los escasos estudios referidos a éstas, efectuados en la región del Cinturón verde La Plata-Buenos Aires, permiten inferir la existencia de enfermedades de las plantas aún no diagnosticadas y la presencia de una mayor diversidad de patógenos no citados en el país, que estarían afectando los cultivos de esa región. Frente a este planteo se propuso el presente estudio con los objetivos de actualizar el estado del conocimiento sobre estos géneros y realizar una prospección de las enfermedades presentes en la región, identificando las nuevas patologías y relaciones hospedante-patógeno, y confirmando la presencia de las previamente citadas. A efectos de recopilar, organizar y actualizar la información referida a especies fitopatógenas de estos géneros en la Argentina, se efectuó una exhaustiva revisión de antecedentes desde finales del siglo XIX hasta el año 2014.
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