\r\n\t\t\t\r\n\t\t\t Figura 1. La principal causa de inducción de defensas en la planta es el ataque de patógenos.<\/strong><\/span><\/p>\r\n\r\n\t\t\tFuente: Intagri.<\/span><\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t<\/tbody>\r\n<\/table>\r\n<\/div>\r\n<\/div>\r\n\r\n\r\n <\/div>\r\n\r\n  \r\n \r\n\t\r\n\t\t\r\n\t\t\t\r\n\t\t\t Figura 2. Respuesta y mecanismos de defensa de plantas ante situaciones bióticas y abióticas adversas.<\/strong><\/span><\/p>\r\n\r\n\t\t\tFuente: Damián, G. J. 2017.<\/span><\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t<\/tbody>\r\n<\/table>\r\n<\/div>\r\n\r\n <\/div>\r\n<\/div>\r\n\r\n \r\n \r\n ¿Qué son los elicitores? <\/strong><\/span><\/h4>\r\n\r\nLos elicitores son moléculas capaces de inducir cualquier tipo de defensa en la planta y son producidos por agentes estresantes bióticos y abióticos. Se puede decir que la aplicación de un elicitor actúa en la planta con el mismo principio de la vacunación; se activa el metabolismo de la planta y se hace más resistente en posteriores ataques que generan estrés. El uso de los elicitores ha crecido por los beneficios que se desencadenan al utilizarlos en los cultivos, pues actúan generalmente en forma de precursores de metabolitos secundarios como las fitoalexinas; estos metabolitos secundarios impiden o retardan la entrada del patógeno a las plantas, pero también limitan su actividad en el tejido u órgano que ha sido infectado. Los elicitores son sustancias naturales o minerales que al ser aplicadas en las plantas de forma preventiva ayudan a reducir o evitar daños producidos por enfermedades, plagas o factores abióticos adversos.<\/p>\r\n\r\n Clasificación de los elicitores <\/strong><\/span><\/h4>\r\n\r\nLos elicitores son muy variados a pesar de que su fin es el mismo, y tienen dos clasificaciones con base en sus características:<\/p>\r\n\r\n a) Según su origen:<\/span><\/h4>\r\n\r\n\r\n\t- Endógenos (endoelicitores): se producen u originan naturalmente dentro de la planta al detectar la necesidad de crear defensas.<\/li>\r\n\t
- Exógenos (exoelicitores): son creados por elicitores externos que se aplican para inducir la producción de defensas en la planta previniendo un ataque, por ejemplo: fosetil-Al, fosfito de potasio, silicio, etc.<\/li>\r\n<\/ul>\r\n\r\n
b) Según su naturaleza:<\/span><\/h4>\r\n\r\n\r\n\t- Bióticos: son moléculas de origen patógeno que pueden inducir respuestas de defensa (como acumulación de fitoalexinas o la respuesta hipersensible) en el tejido vegetal. En su mayoría compuestos orgánicos producidos por el agente estresante (carbohidratos, lípidos, enzimas microbianas, ácido salicílico).<\/li>\r\n\t
- Abióticos: generados por factores abióticos como condiciones ambientales (frío, luz UV, metales pesados, detergentes).<\/li>\r\n<\/ul>\r\n\r\n
¿Cómo actúan los elicitores? <\/strong><\/span><\/h4>\r\n\r\nEn la etapa de infección temprana del patógeno, el reconocimiento de éste es muy importante para que la planta logre defenderse. Se supone que el reconocimiento del elicitor por la planta está mediado por receptores específicos en la célula de la planta, los cuales inician los procesos de señalización que activan las defensas de las plantas. Un mecanismo general para la elicitación biótica en plantas puede resumirse sobre la base de la interacción elicitor-receptor. Cuando una planta o cultivo de células vegetales es desafiado por el estimulante se produce una serie de actividades bioquímicas; algunas actividades bioquímicas que se desencadenan con la aplicación de elicitores en la planta son: generación de especies reactivas de oxígeno (ROS), acumulación de proteínas relacionadas con la patogénesis como quitinasas y gluconasas, muerte celular en el sitio de infección (respuesta hipersensible), cambios estructurales en la pared celular (lignificación de la pared celular), activación transcripcional de los correspondientes genes de respuesta de defensa, síntesis de moléculas defensivas de las plantas como los taninos y las fitoalexinas, síntesis de ácidos jasmónicos y salicílicos como mensajeros secundarios, y finalmente la resistencia sistémica adquirida.<\/p>\r\n<\/div>\r\n<\/div>\r\n\r\n \r\n \r\n \r\n\t\r\n\t\t\r\n\t\t\t\r\n\t\t\t Cuadro 1. Sustancias elicitoras y su efecto de control en cultivos específicos.<\/strong><\/span><\/p>\r\n\r\n\t\t\tFuente: Wiesel et al.<\/em>, 2014.<\/span><\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Elicitor<\/strong><\/p>\r\n\t\t\t<\/td>\r\n\t\t\t\r\n\t\t\t Efectivo contra<\/strong><\/p>\r\n\t\t\t<\/td>\r\n\t\t\t\r\n\t\t\t Cultivo (Referencia)<\/strong><\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Oligogalacturónidos<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Botrytis cinerea, Blumeria graminis<\/em><\/p>\r\n\t\t\t<\/td>\r\n\t\t\t\r\n\t\t\t Varios (Aziz et al<\/em>., 2007; Randoux et al<\/em>., 2010; Galletti et al., 2011)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Extracto de Reynoutria sachalinensis<\/em><\/p>\r\n\t\t\t<\/td>\r\n\t\t\t\r\n\t\t\t Botrytis cinerea, Leveillula<\/em> \r\n\t\t\ttaurica<\/em><\/p>\r\n\t\t\t<\/td>\r\n\t\t\t\r\n\t\t\t Pepino, tomate (Daayf et al<\/em>., 1997, 2000; \r\n\t\t\tKonstantinidou-Doltsinis et al<\/em>., 2006)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Fructooligosacárido de Arctium lappa<\/em><\/p>\r\n\t\t\t<\/td>\r\n\t\t\t\r\n\t\t\t Colletotrichum lagenarium,<\/em> \r\n\t\t\tBotrytis cinerea, <\/em>TMV<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t\r\n\t\t\t Pepino, tomate, tabaco (Wang et al<\/em>., 2009; Guo et al<\/em>., 2012)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Elicitor peptídico 1<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Cochliobolis heterostrophus<\/em>, \r\n\t\t\tColletotrichum graminicola<\/em><\/p>\r\n\t\t\t<\/td>\r\n\t\t\t\r\n\t\t\t Maíz (Huffaker et al<\/em>., 2011)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Carragenanos<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Sclerotinia sclerotiorum, <\/em>TMV<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t\r\n\t\t\t A. thaliana, tabaco (Sangha et al.<\/em>, 2010; Vera et al<\/em>., 2011)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Fucanos<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t TMV<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Tabaco (Vera et al<\/em>., 2011)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Ulvans<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Varios<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Varios (Jaulneau et al<\/em>., 2011; Vera et al<\/em>., 2011)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Laminarina<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Erwinia carotovora,<\/em> \r\n\t\t\tPlasmopara viticola, Botrytis<\/em> \r\n\t\t\tcinerea<\/em>, Fusarium solani<\/em><\/p>\r\n\t\t\t<\/td>\r\n\t\t\t\r\n\t\t\t Frijol, uva de vino, tabaco (Craigie, 2011; Vera et al<\/em>., 2011)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t<\/tbody>\r\n<\/table>\r\n<\/div>\r\n<\/div>\r\n\r\n\r\n \r\n Compuestos comerciales inductores de la defensa de las plantas<\/strong><\/span><\/h4>\r\n\r\nFosfitos de potasio<\/strong>. Inductores de síntesis de fitoalexinas al estimular enzima PAL (Fosfo-amino-liasa), clave en la producción de compuestos tipo fenoles (resveratrol), ligninas, suberinas y compuestos derivados del ácido cinámico; todos estos son mecanismos de defensa de la planta. Cuentan con alta movilidad dentro de la planta y las fitoalexinas inducidas tienen efecto sobre hongos de la familia de los oomicetos. Su uso se recomienda cuando no existan deficiencias de fósforo, ya que puede presentar efectos contraproducentes.<\/p>\r\n\r\nGlutation<\/strong>. Es formado por la unión de ácido glutámico, cisteína y glicina. Actúa como reductor de reacciones metabólicas, es un componente de la resistencia de plantas a enfermedades y tiene acción desintoxicante en las células.<\/p>\r\n\r\nOligosacarinas<\/strong>. Cadenas de glucósidos producidas por la desintegración de paredes celulares por actividad de enzimas de patógenos. Estas moléculas activan genes de resistencia por hipersensibilidad que inducen la producción de compuestos de protección.<\/p>\r\n\r\nSaponinas<\/strong>. Se encuentra en el tejido de soporte de las plantas, cumple función de protección desintegrando la membrana del organismo patógeno que cause estrés.<\/p>\r\n\r\nLa mayoría de los elicitores desencadenan la liberación de compuestos de defensa pero además se descubren que tienen efecto de control de enfermedades específicas como Fusarium spp.<\/em>, Phytophthora spp.<\/em>, Alternaria spp.<\/em> (Cuadro 2).<\/p>\r\n<\/div>\r\n<\/div>\r\n\r\n\r\n \r\n \r\n\t\r\n\t\t\r\n\t\t\t\r\n\t\t\t Cuadro 2. Listado de elicitores y su efecto en diferentes plantas.<\/strong><\/span><\/p>\r\n\r\n\t\t\tFuente: Thakur y Singh, 2012.<\/span><\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Planta<\/strong><\/p>\r\n\t\t\t<\/td>\r\n\t\t\t\r\n\t\t\t Elicitor<\/strong><\/p>\r\n\t\t\t<\/td>\r\n\t\t\t\r\n\t\t\t Efecto (Referencia)<\/strong><\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Tomate<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Ácido Salicílico<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Disminuye la susceptibilidad a patógenos y estrés abiótico. Induce síntesis de proteínas de estrés, que incrementan resistencia al frío. (Shirasu et al.<\/em>, 1997; García et al.<\/em>, 2002)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Ácido Oligogalacturórico<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Reducen apertura estomática; acelera el cierre de estomas. (Lee et al.<\/em>, 1999)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Quitosano<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t El quitosano inhibe apertura estomática inducida por luz y produce mayor resistencia a Fusarium oxysporum<\/em> y Phytophthora capsici<\/em>. (Ortega et al.<\/em>, 2003; Ortega et al.<\/em>, 2007)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Genes Antivirulentos (AVR)<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Activación de respuesta hipersensitiva. (Leach and White, 1996).<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t | \r\n\t\t\t\r\n\t\t\t Colza<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Jasmonato de Metilo<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Acumulación de indolil glucosinolatos (repelentes) en las hojas. (Doughty et al.<\/em>, 1995).<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Ácido Salicílico y Óxido Nítrico<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Aumenta actividades de enzimas antioxidantes, mejora el contenido de clorofila y disminuye el nivel de peroxidación de lípidos. (Kazemi et al.<\/em>, 2010).<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Arroz<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Benzotiazol<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Protege contra mildiu polvoso afectando el ciclo de vida del patógeno. (Görlach et al.<\/em>, 1996).<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Col<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Benzotiazol<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Induce síntesis de quitinasa y resistencia al virus de necrosis del tabaco y mildiu velloso en plantas de semillero y de 30 días. (Godard et al.<\/em>, 1999).<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Ácido salicílico, Quitosano, Jasmonato de metilo.<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t El ác. Salicílico y quitosano inducen incremento en contenido de vitamina C. Tratamientos con Jasmonato de metilo y á. salicílico incrementaron la concentración de flavonoides. (Pérez et al.<\/em>, 2011)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Frijol<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Ácido Salicilico y Jasmonato de Metilo<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Controlan infestaciones de ácaros, incrementa crecimiento y rendimiento. (Farouk y Osman, 2011)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t | \r\n\t\t\t\r\n\t\t\t Soya<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Benzotiazol<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Disminución de la decoloración del tallo, aumento de la germinación, pigmentos fotosintéticos, compuestos fenólicos y flavonoides. (Nafie y Mazen, 2008)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t | \r\n\t\t\t\r\n\t\t\t Benzotiazol y ácido húmico<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Reduce Damping-off<\/em> y marchitez, e incrementa parámetros de crecimiento. (Abdel et al.<\/em>, 2011)<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t\t\r\n\t\t\t\r\n\t\t\t Remolacha<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Benzotiazol<\/p>\r\n\t\t\t<\/td>\r\n\t\t\t | \r\n\t\t\t Induce síntesis de quitinasa y provee resistencia al virus de necrosis del tabaco. (Burketová et al., 1999).<\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t<\/tbody>\r\n<\/table>\r\n<\/div>\r\n<\/div>\r\n\r\n \r\n \r\n Cita correcta de este artículo<\/strong><\/span><\/p>\r\n\r\nINTAGRI. 2017. La Inducción de Defensa en las Plantas a través de Elicitores. Serie Fitosanidad Núm. 92. Artículos Técnicos de INTAGRI. México. 6 p.<\/p>\r\n\r\n Fuentes consultadas:<\/strong><\/span><\/p>\r\n\r\nDamián, G. J. 2017. Inducción de las Defensas en las Plantas a través de Elicitores. Diplomado Internacional de Bioestimulación de Cultivos Intensivos. Intagri. Gto. México. <\/p>\r\n\r\n Wiesel, L.; Newton, A.C.; Elliot, I.; Boooty, D.; Gilroy, E.M.; Birch, P.J.R.; Hein, I. 2014. Molecular Effects of Resistance Elicitors from Biological Origen and Their Potencial for Crop Protection. Frontiers in Plant Science, 5:1-13.<\/p>\r\n\r\n Thakur, M.; Singh, S. B. 2013. Role of Elicitors in Inducing Resistance in Plants against Pathogen Infection: A Review. ISRN Biochemistry, 2013: 1-10.<\/p>\r\n\r\n Angelova, Z.; Georgiev, S.; Roos, W. 2006. Elicitacion of Plants. 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El uso de los elicitores ha crecido por los beneficios que se desencadenan al utilizarlos en los cultivos, pues actúan generalmente en forma de precursores de metabolitos secundarios como las fitoalexinas; estos metabolitos secundarios impiden o retardan la entrada del patógeno a las plantas.","fb_desc":"","thumb":"\/assets\/images\/articulos\/categoria1\/Fitosanidad\/art419-induccion-de-defensa-de-plantas\/Elicitores%20Fitoalexinas.jpg","fb_thumb":"","category":8,"related":"","keywords":"Elicitores,Elicitors,Inductores de Resistencia,Fitoalexinas,Phytoalexins","pdfLink":"Articulos\/Fitosanidad\/92. La Induccion de Defensa en las Plantas a traves de Elicitores.pdf","text":" \r\n \r\n Las plantas continuamente están sometidas a situaciones bióticas y abióticas adversas, y ante cualquier ataque de patógenos o condición de estrés las plantas cuentan con mecanismos químicos de defensa<\/a>. Dentro de los mecanismos de defensa se pueden enunciar dos tipos: 1) los pasivos y 2) los activos. Los primeros están referidos a las defensas físicas de las plantas como cutículas, tricomas y ceras, así como a las barreras químicas, que consisten en la síntesis de sustancias químicas por parte de la planta antes de cualquier infección por patógenos; tales sustancias son saponinas, alcaloides, proteínas antifúngicas o enzimas inhibidoras. Por otra parte, los mecanismos activos tienen que ver generalmente con la producción endógena o aplicación exógena de compuestos conocidos como “elicitores”, cuyo objetivo de estos es fungir como activadores de reacciones defensivas, es decir, inducir la producción de fitoalexinas o estimular cualquier mecanismo de defensa de la planta para protegerse. Estos inductores son capaces de promover diferentes modos de defensa de la planta, como: Resistencia Sistémica Adquirida (relacionada al ácido salicílico y proteínas PR), Resistencia Sistémica Inducida (activada por cepas bacterianas de rizobacterias saprofitas) y Resistencia Local Adquirida (desencadenada por la respuesta hipersensible de la planta y la producción de fitoalexinas) (Figura 2<\/strong>).<\/p>\r\n\r\n¿Qué son las fitoalexinas? <\/strong><\/span><\/h4>\r\n\r\nLas fitoalexinas son producidas después de una infección por la interacción planta (hospedero) y patógeno (huésped) para frenar la infección; tienen como características que no se detectan antes de la infección, se sintetizan aproximadamente de 1 a 8 horas después del ataque, solo se producen alrededor del punto de infección y son tóxicas a un amplio espectro de hongos y bacterias. Las sustancias que estimulan las síntesis de fitoalexinas se denominan elicitores.<\/p>\r\n<\/div>\r\n\r\n  \r\n \r\n\t\r\n\t\t\r\n\t\t\t\r\n\t\t\t Figura 1. La principal causa de inducción de defensas en la planta es el ataque de patógenos.<\/strong><\/span><\/p>\r\n\r\n\t\t\tFuente: Intagri.<\/span><\/p>\r\n\t\t\t<\/td>\r\n\t\t<\/tr>\r\n\t<\/tbody>\r\n<\/table>\r\n<\/div>\r\n<\/div>\r\n\r\n\r\n <\/div>\r\n\r\n  \r\n \r\n\t\r\n\t\t\r\n\t\t\t\r\n\t\t\t
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