Pudrición de raíz y tallo de Soja (Phytophthora sojae)

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Condición fitosanitaria: Presente ampliamente distribuida

Grupo de cultivos: Oleaginosas

Especie hospedante: Soja (Glycine max)

Rango de hospedantes: específico / estrecho. La soja es el único hospedante económicamente importante. También se han reportado varias especies de lupinos (Lupinus spp.) como hospedantes (Tyler, 2006).

Epidemiología: monocíclica, subaguda.

Etiología: Pseudohongo. Necrotrófico (breve fase inicial hemibiotrófica entre 24 a 36 hpi)

Agente causal: Phytophthora sojae  Kauffmann & Gerdemann

Taxonomía: Eukaryota > Stramenopiles > Oomycetes > Peronosporales > Peronosporaceae > Phytophthora

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Antecedentes

Esta enfermedad fue una de las primeras patologías detectadas en el cultivo de soja, luego de la podredumbre de tallos y vainas por Sclerotinia sclerotiorum y la mancha marrón por Septoria glycines.

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Síntomas

El síntoma típico de la enfermedad está constituido por la decoloración parda del tallo desde la superficie del suelo que contrasta con los tejidos verdes superiores. La pudrición de raíces y de la base del tallo conduce finalmente al marchitamiento de las plantas. El patógeno infecta en cualquier etapa del cultivo. Puede causar tizón en pre- y post-emergencia (ver tizón de plántulas). Los síntomas generales, incluyen amarillamiento de hojas y marchitamiento de las plantas. Los síntomas en plantas adultas antes del marchitamiento consisten en el oscurecimiento de la base del tallo que avanza por las ramificaciones hasta el quinto o sexto entrenudo y contrasta con los tejidos verdes superiores aún sanos. Las raíces y el tejido vascular son invadidos por el patógeno. Finalmente se produce la muerte de la planta y las hojas quedan adheridas al tallo. Cultivares tolerantes pueden mostrar pudrición del sistema radical, pero sobreviven.

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Condiciones ambientales predisponentes para el establecimiento de la enfermedad, Epidemiología

El patógeno es un oomycete, y por lo tanto forma esporas flageladas (zoosporas) que se dispersan fácilmente en medio acuoso. Este oomycete tiene la capacidad de persistir en el suelo y en los rastrojos como oosporas. Las oosporas permanecen latentes durante años, ya que las esporas producidas en la misma estación generalmente deben acumular un período de suelo frío antes de que ocurra la germinación.

Las condiciones ambientales predisponenetes son suelos húmedos y fríos, pobremente drenados, pesados (con altos contenidos de arcilla), presencia de rastrojo infectado y drenaje deficiente, predisponen al desarrollo de la enfermedad, la cual se manifiesta también con mayor intensidad en áreas bajas de los lotes. El desarrollo de epidemias se ve favorecido por la acumulación de agua en el suelo ya que facilita la dispersión de las zoosporas, por lo cual los daños son más severos en suelos compactos, en las cabeceras de los lotes y en años lluviosos. La presencia de rastrojos infestados del cultivo anterior aumenta la densidad de inóculo del patógeno ya que las oosporas quedan protegidas por los tejidos muertos.

En condiciones de suelo casi saturado, las oosporas preacondicionadas germinan para formar micelio que luego produce zoosporangios. Los zoosporangios maduran rápidamente y cuando el suelo se satura, liberan zoosporas para infectar las raíces del hospedante. Las zoosporas son negativamente geotácticas y son atraídas por exudados de raíces como daidzeina y genisteína  (isofalvonoide). Una vez liberadas, las zoosporas nadan hacia las raíces de la planta hospedante. Una vez que entran en contacto con una raíz, las zoosporas se enquistan, pierden sus flagelos y forman una pared celular. Luego, germinan para formar un tubo germinativo y penetran en la epidermis de la raíz a través de la formación de un apresorio.

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Manejo de la enfermedad

P. sojae es un patógeno de alta especificidad, si bien la soja no es el único hospedante, no ataca otros cultivos extensivos de verano en la Argentina. La enfermedad es fácilmente controlable a través de la resistencia genética que es conferida por genes mayores Rps. Durante muchos años se observó baja variabilidad en virulencia ya que prevaleció la raza 1 (Barreto et al, 1995, 1998). Sin embargo, la liberación de cultivares resistentes indujo la aparición o incremento de frecuencia de nuevas razas lo cual dificulta el manejo de la enfermedad.

Desde hace varios años, se han encontrado fuentes de resistencia a P. sojae (Dorrance y Schmitthenner, 2000; Yang et al., 2020; Bolaños-Carriel et al., 2021). En las últimas décadas, se han mapeado unas pocas docenas de loci / alelos de resistencias (Rps) en varias regiones genómicas de la soja enriquecidas con genes que codifican proteínas receptoras de resistencia específicas (proteínas de sitio de unión a nucleótidos de repetición rica en leucina (NBS-LRR o NLR) en base al ensamblaje del genoma de referencia de soja v2.0 (soybase.org) (Sahoo et al., 2017; Zhong et al., 2018; Matthiesen et al., 2016; Jiang et al., 2020; Matthiesen et al., 2021). Sin embargo, debido a la gran variabilidad genética en las poblaciones del patógeno en constante evolución (Abney et al., 1997; Yang et al., 2019), muchos de esos loci / alelos se han vuelto ineficaces para los aislados del patógeno recientemente evolucionados y se han aislado y/o validado funcionalmente pocos loci/alelos de Rps (Dong et al., 2011; Yan y Nelson, 2019; Yang et al., 2019). Por lo tanto, la introgresión/piramidación efectiva de genes Rps específicos en cultivares de soja de élite para lograr una resistencia sostenida sigue siendo un desafío. Recientemente, Wang et al. (2021) encontraron que el gen de resistencia Rps11 confiere resistencia de amplio espectro a P. sojae; y Sahoo et al. (2021) encontraron que dos genes Rps ( Rps12 and Rps13) funcionales estrechamente ligados con una especificidad de raza distinta proporcionan una resistencia de amplio espectro en la soja.

Para el manejo de este patosistema es necesario integrar varias herramientas de manejo, como uso de cultivares resistentes, rotaciones (2 a 3 años con gramíneas) y tratamiento de semillas con metalaxil. Mejorar el drenaje si es posible. P. sojae tiene alta variabilidad patogénica (razas) actualmente en nuestro país, por lo cual los cultivares que resistentes a la raza 1, que es la más común, pueden no serlo para las que se encuentren en el lote afectado.

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Phytophthora Root and Stem Rot of Soybean. Crop Protection Network. Iowa State University Integrated Pest Management Program.

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