Tizón del arroz (Pyricularia oryzae / Magnaporthe oryzae)

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Condición fitosanitaria: Presente

Grupo de cultivos: Cereales

Especie hospedante: Arroz (Oryza sativa)

Rango de hospedantes: M. oryzae tiene un rango amplio de hospedantes, pero los aislados individuales tienen rangos de hospedantes limitados y se dividen en diferentes patotipos según su compatibilidad (Chung et al., 2020).

Epidemiología: policíclica, subaguda.

Etiología: Hongo. Hemibiotrófico

Agente causal:  Magnaporthe oryzae (anamorph: Pyricularia oryzae  Cavara, 1892)

Taxonomía: Eukaryota > Fungi > Dikarya > Ascomycota > Pezizomycotina > Sordariomycetes >  Magnaporthales > Magnaporthe

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Sintomatología

Pyricularia oryzae ataca hojas, tallos, inflorescencias y ocasionalmente al grano. Los momentos, en que la planta de arroz es más susceptible, son el estado de plántula y durante la floración, incluso las raíces pueden infectarse. Sin embargo, el síntoma más común y que sirve como diagnóstico, son lesiones en forma de diamante que ocurre en las hojas, mientras que las lesiones en las vainas son relativamente raras.

Hojas: los síntomas en las hojas pueden variar de acuerdo con las condiciones ambientales, la edad de la planta y los niveles de resistencia de los cultivares hospedantes. En cultivares susceptibles, las lesiones pueden aparecer inicialmente con un color verde grisáceo, con un borde verde más oscuro y se expanden rápidamente a varios centímetros de longitud. En cultivares susceptibles, las lesiones más viejas a menudo adquieren un color tostado claro con bordes necróticos. En cultivares resistentes, las lesiones suelen ser pequeñas en tamaño (1-2 mm) y de color café a marrón oscuro.

Unión de la hoja y la vaina del tallo: los síntomas de infección consisten en un área general de necrosis en la unión de los dos tejidos. Este tipo de infecciones pueden matar toda la hoja y extenderse unos pocos milímetros dentro y alrededor de la cubierta. El hongo puede producir esporas en estas lesiones.

Cuellos y panículas: el cuello de la planta de arroz se refiere a la porción del tallo que se eleva sobre las hojas y sostiene la cabeza o la panícula de la semilla. Los cuellos a menudo se infectan en el nódulo y la infección conduce a una afección llamada cuello podrido o explosión del cuello. La infección de los cuellos puede ser muy destructiva, causando que las semillas no se llenen, o haciendo que toda la panícula se caiga como si estuviera podrida. El hongo también puede infectar las panículas a medida que se forman las semillas. Las lesiones se pueden encontrar en las ramas de la panícula, espinas y espiguillas. Las lesiones a menudo son decoloraciones de color marrón grisáceo de las ramas de la panícula y, con el tiempo, las ramas pueden romperse en la lesión.

Cuando la infección se inicia en un nudo, éste toma un color grisáceo y la parte superior de la planta se seca. Esta zona suele ser un punto de rotura de la caña.

Semillas: el hongo a menudo se ha aislado de los pedicelos de las semillas. Las semillas no se producen cuando los pedicelos se infectan, una condición llamada cegamiento. Los síntomas de la explosión del arroz en las semillas mismas consisten en manchas marrones, y ocasionalmente, la lesión clásica en forma de diamante de las hojas. El proceso y el tiempo durante el cual ocurre la infección de las semillas por las esporas del patógeno no se ha descrito completamente, pero la información reciente muestra que el hongo puede infectar las semillas infectando las flores mientras maduran en semillas, y se cree que este es la principal manera en que se desarrolla la infección de semilla.

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Hospedantes

Afecta un amplio espectro de especies de las Gramíneas, pero el principal daño económico lo provoca en arroz.

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Condiciones predisponentes

Alta humedad relativa (90 – 92 %), gran intensidad lumínica, periodos largos de fotoperiodismo, tiempo ventoso, temperaturas nocturnas rondando los 20 ºC alternando con temperaturas diurnas entre 30 – 35 ºC.

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Ciclo de la enfermedad y epidemiología

M. oryzae produce esporas sexuales (ascosporas) en estructuras llamadas asci (ascos), y se clasifica en la familia Magnaporthaceae recientemente erigida. Los asci se encuentran dentro de estructuras especializadas llamadas peritecios. El micelio de M. oryzae está septado y los núcleos dentro del micelio y las esporas de este hongo son haploides.

Como ascomicete, produce ascosporas hialinas, de forma fusiforme (con forma de huso con extremos decrecientes) con tres tabiques. Los asci son unitunicados. Se considera que este hongo es heterotálico con un sistema de acoplamiento bipolar (apareamiento controlado por dos alelos diferentes en un único locus) con genes adicionales que controlan el ciclo sexual. Con base en datos filogenéticos, moleculares y morfológicos recientes, aislamientos del hongo del arroz y aislamientos estrechamente relacionados de otros pastos como Eragrostris curvula, Eleusine coracana, Lolium perenne y Setaria spp. se describen taxonómicamente como Magnaporthe oryzae, mientras que los aislamientos de Digitaria sanguinalis son distintos y deben describirse como Magnaporthe grisea.

La etapa asexual de Magnaporthe oryzae se describe con el nombre de Pyricularia oryzae (antes llamada P. grisea) y es la forma de espora más común del hongo en Argentina. Estas esporas, llamadas conidios, se producen abundantemente en lesiones y en cultivo en tallos especializados, llamados conidióforos. Los conidios generalmente son tricelulares y se producen en el ápice de un conidióforo. Las colonias esporuladas en placas de agar pueden adquirir una apariencia grisácea y velluda.

En condiciones favorables, el hongo esporula en el centro de las lesiones en cultivares susceptibles. También puede esporular en lesiones de semillas. Raramente esporula en los cultivares más resistentes. Las esporas se producen en hojas, collar, panículas y semillas infectadas, en conidióforos que se extienden más allá de las superficies lesionadas; los conidióforos y las esporas en masa pueden dar a las lesiones un aspecto gris polvoriento. Los conidios se producen después de varias horas de alta humedad y se liberan más fácilmente cerca del mediodía, especialmente en condiciones de viento. El viento puede llevar los conidios a distancias considerables.

La infección del arroz ocurre cuando los conidios se depositan en los tejidos de arroz y germinan al producir un tubo germinativo y un apresorio. El apresorio es una estructura melanizada, y de ella se desarrolla una clavija de infección que penetra el tejido. El hongo penetra directamente por la cutícula. Después de la penetración, la hifa de infección primaria crece rápidamente y se ramifica dentro de los tejidos susceptibles. El crecimiento dentro de los tejidos de los cultivares resistentes a menudo se inhibe.

En general, el tizón del arroz se ve favorecido por temperaturas moderadas (24°C) y períodos de alta humedad que son de 12 horas o más, condiciones fácilmente alcanzables en campos de arroz inundados.

Las fuentes de esporas invernantes que comprenden el inóculo primario consisten en hierbas, plantas voluntarias, rastrojos infestados y semillas infestadas en la superficie del suelo después de la siembra mecánica. Las semillas infestadas que quedan en la superficie del suelo pueden producir fácilmente esporas de P. oryzae durante más de varias semanas después de la siembra, mucho después de que hayan surgido las plántulas. En invernadero, el hongo también esporula sobre los coleópticos muertos o moribundos de plantas que crecen a partir de semillas infectadas.

Las esporas producidas como inóculo primario en los tejidos que invernan, producen las infecciones iniciales en plántulas jóvenes, al depositarse en las hojas, germinando e invadiendo los tejidos de las hojas. La gravedad de la enfermedad a menudo se correlaciona con la cantidad de material infestado (densidad de inóculo). Las lesiones en las plántulas jóvenes aparecen unos días después de la infección. Estas lesiones secundarias producen más esporas y estas esporas se diseminan fácilmente por el viento a los tejidos de hojas sanos cercanos. La esporulación ocurre cuando la humedad relativa alcanza al 100 %.

Los ciclos secundarios pueden repetirse muchas veces durante la temporada de crecimiento, con el potencial de generar una gran cantidad de infecciones dentro de la estación de cultivo. El número de ciclos y el número de esporas que se producen en cada lesión individual pueden verse influenciados por muchos factores, que incluyen la temperatura, la lluvia, la profundidad del agua en el arrozal, la cantidad de nitrógeno utilizada para fertilizar el arroz y la nivel de resistencia genética en el cultivar que está infectado.

En general, la enfermedad en las hojas es más severa cuando las temperaturas diarias son moderadas, y cuando el arroz está sobre-fertilizado, o si se cultiva en aguas de inundación por debajo de las profundidades recomendadas. Bajo estas condiciones propicias para la enfermedad, se han registrado muy altas incidencias de la enfermedad en cultivares susceptibles, en comparación con cultivares genéticamente resistentes al hongo.

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Manejo Integrado de la enfermedad

* Rotación de cultivos

* Usar semilla libre de enfermedad

* No excederse en la fertilización nitrogenada

* Uso de fungicidas: los momentos de tratamiento que se consideran más eficaces son formación de espiga e inicio floración.

* Control biológico: se han reportado casos satisfactorios de biocontrol del patógeno (Sha et al., 2020; Zhu et al., 2021)

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Videos

Rice Diseases: Rice Blast

What can we do to actively manage wheat blast?

Septin-Dependent Assembly of the Exocyst Is Essential for Plant Infection by Magnaporthe oryzae

Mitosis in the appressorium and invasive hypha of the rice blast fungus (www.khanglab.org)

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