Fisiología de la Nutrición Mineral Ing. M Sc. Hilvio Castillo
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2.1 General
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Un nutriente esencial es: • IMPRESCINDIBLE: sin su presencia la planta
no se desarrolla normalmente o no completa su ciclo de vida • ESPECÍFICO: no puede ser reemplazado por
otro elemento
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2.1 General Por lo general hay una gran discrepancia entre: •La concentración de nutrientes minerales en el suelo o solución nutritiva, y •El requerimiento vegetal de nutrientes nu trientes minerales. Además, el suelo y también en algunos casos las soluciones nutritivas pueden contener altas concentraciones de elementos minerales no necesarios para el crecimiento vegetal. Por lo tanto, los mecanismos por los cuales las plantas toman los nutrientes deben ser selectivos. Esta selectividad puede ser bien demostrada particularmente en células de algas (Tabla 2.1), donde las soluciones externas e internas (savia celular) están separadas por solo dos membranas: la membrana
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Aunque usualmente menos dramática, también es un característica típica en plantas superiores la selectividad en la toma iónica . Cuando las plantas son cultivadas en volúmenes limitados de solución nutritiva, la concentración externa cambia en pocos días (Tabla 2.2). Las concentraciones de potasio, fosfato y nitrato disminuyen marcadamente, mientras que pueden aumentar las de sodio y sulfato, indicando que el agua es tomada más rápido que cualquiera de estos dos iones. Entre las dos especies vegetales (maíz y fríjol) difieren las tasas de toma, especialmente para el potasio y calcio . La concentración iónica en la savia radical exprimida es generalmente mayor que en la solución nutritiva; esto es mas evidente en el caso del potasio, nitrato y
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Los resultados obtenidos en ambas plantas inferiores y superiores demuestran que la toma de iones se caracteriza por lo siguiente:
Selectividad. Ciertos elementos minerales son tomados preferentemente, mientras que otros son en contra discriminados ó casi excluidos Acumulación. La concentración de elementos minerales puede ser
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Fig. 2.1 Sección transversal de dos células rizodérmicas en una raíz de maíz. V, vacuola; C, citoplasma; W, pared celular; E, solución externa. (Cortesía de C. Hecht-Buchholz)
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Las paredes celulares primarias consisten de una red de celulosa, hemicelulosa, (incluyendo pectinas) y glicoproteínas, las ultimas pueden representar entre 5 y 10% del peso seco de las paredes celulares. Esta red contiene poros, los llamados espacios intermicelares e interfibrilares, que difieren en tamaño. Se ha calculado para células de pelos radicales de rábano un diámetro máximo de 3.5 – 3.8 nm (35-38 Å); los valores máximos para poros en paredes celulares vegetales están en el rango de 5.0 nm. En comparación, los iones hidratados como K+ y Ca 2+ como se muestra abajo son pequeños estando en el orden de solo 10 – 20% del
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En contraste a los nutrientes minerales y solutos orgánicos de bajo peso molecular, los solutos de alto peso molecular (e.g., quelatos metálicos, ácidos fúlvicos, y toxinas) o virus y otros patógenos cualquiera son severamente limitados o evitados por el diámetro de los poros de entrar al espacio libre de la células radicales . En esta red, una proporción variable de pectinas consisten de ácido poligalacturónico originado principalmente de la lamela media. Por consiguiente ambos en raíces y en el continuo de pared celular de otros tejidos vegetales, el llamado apoplasto, los grupos
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Fig. 2.2 Diagrama esquemático del sistema de poros del espacio libre aparente. DFS, espacio libre de Donan; WFS, espacio libre acuoso, gracias al potencial electroquímico generado por iones no difusibles se supera el equilibrio a uno de los lados de una membrana.
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La distribución iónica dentro del DFS se caracteriza por la típica
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Por lo general, la CEC de especies dicotiledóneas es mucho mayor que en especies monocotiledóneas. monoc otiledóneas. La CEC efectiva disminuye como
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La importancia del ligamiento catiónico en el AFS para la toma y subsiguiente transporte al vástago v ástago también es indicada por
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Los ácidos monocarboxílicos no disociados pueden incrementar el agujereado de la membrana al cambiar la composición de ácidos grasos de la membrana, particularmente al disminuir la proporción de ácidos grasos poliinsaturados como el ácido linolénico. Es de considerable importancia ecológica el efecto de los ácidos monocarboxílicos sobre la permeabilidad de la membrana radical, ya que estos ácidos se acumulan en suelos inundados (Sección 16.4). Como se muestra en la Fig. 2.5, estas especies tóxicas de oxigeno son cualquiera radicales como el superóxido (O 2.-) ó hidroxilo (OH.), ó la molécula peróxido de hidrogeno (H 2O2). Todas se forman en varias reacciones y procesos metabólicos donde el oxigeno este involucrado, por ejemplo en la fotosíntesis y en la respiración, incluyendo la oxidación de NADPH ó NADH en la interfase membrana plasmática –pared celular. La toxicidad por especies activadas de oxigeno y sus derivados se causa, por ejemplo por la oxidación de grupos tioles (-SH) de enzimas y la peroxidación de los ácidos grasos
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Fig. 2.5 Modelo de generación de, y peroxidación lipídica membrana por, radicales de oxígeno y peróxido de hidrógeno, y sistemas de secuestro y detoxificación. (I. Cakmak, no publicado.)
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La nutrición mineral vegetal puede afectar en varios niveles ambos, la generación de especies tóxicas de oxigeno de peróxido de hidrogeno, y los mecanismos para su detoxificación. Esto puede resumirse así: 1. Como un componente de enzimas detoxificantes (e.g., zinc, cobre, manganeso, o hierro en las superóxido dismutasas; hierro en peroxidasas y catalasas); 2. Por acumulación de precursores para la formación de radicales (e.g., fenoles y quinonas) bajo deficiencia nutricional (e.g., deficiencia de boro); 3. A través de la disminución en la actividad demanda (i.e., la demanda) y la acumulación de fotosintatos y correspondientemente elevados niveles de especies tóxicas de oxigeno en hojas fuente bajo deficiencia de nutrientes minerales (e.g., potasio y magnesio).
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2.4 Transporte de solutos a través de membranas
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2.4 Transporte Transporte de solutos a través de membranas
2.4.1 Transporte y demanda energética. 2.4.1 Transporte
2.4.2 Transporte Transporte activo y pasivo: pasivo: bombas bombas electrogénicas, carriers, canales iónicos. 2.4.3
La cinética del transporte.
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2.4.1 Transporte y
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2.4.1 Transporte y demanda energética Las membranas intactas son barreras efectivas al paso de iones y moléculas no cargadas. Por otro lado, ellas también son centros de selectividad y transporte contra el gradiente de concentraci ón de los solutos. solutos. En el experimento registrado en la Tabla 2.2, por ejemplo, la
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Tabla 2.2 2. 2 Cambios en la concentración iónica en la solución (nutritiva) externa y en la savia radical exprimida en maíz y fríjol
Concentración externa (m м) Después de 4 días a
Concentración en la savia radical exprimida (mм)
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Fig. 2.6 (A) Toma del ión potasio (influjo) y (B) actividad ATPasa (ATP → ADP + Pi) en raíces aisladas de diferentes especies vegetales. Clave: , cebada; , avena; , trigo; , maíz. (Después de Fisher et al .,., 1970.)
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Se propuso en este proceso un acoplamiento directo del transporte selectivo de iones mediado por carriers y el consumo de fosfatos ricos en energía en forma de ATP. En este modelo se suministro ATP vía respiración (fosforilación oxidativa en la mitocondria; Sección 8.4.3) y se requirió para la activación del carrier, para el ligamiento del ión al carrier, para el transporte membranal del complejo carrier –ión, ó para la liberación liberación del ión del carrier en la superficie interna de la membrana. Fisher et al (1970)
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Es considerable la demanda energética para la toma iónica radical, especialmente durante el rápido crecimiento vegetativo (Tabla
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Estos cálculos al nivel de planta entera sobre la demanda de ATP para la toma iónica radical tienen que interpretarse con cuidado respecto a la demanda energética para el transporte membranal
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El transporte de solutos a través de las membranas no es necesariamente un proceso activo. Los solutos pueden estar mas concentrados en un lado de la membrana (i.e., ellos puede poseer mas
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En el modelo mostrado en la Fig. 2.9 la H +-ATPasa de la membrana plasmática juega un rol clave en ambos la regulación del pH citoplásmico y en la fuerza motriz para la toma catiónica y aniónica. Se considera por lo tanto como la “enzima maestra” . Se ha hecho considerable progreso en el entendimiento de cómo es regulada la actividad de esta enzima, también a nivel genético. La actividad de la H +-ATPasa de la membrana plasmática es particularmente alta en pelos radicales donde esta consume tanto como 25-50% del ATP ATP celular. cel ular. Por consiguiente, bajos niveles radicales de carbohidratos no solo disminuyen la extrusión neta de protones hacia el medio externo sino también disminuye el pH citosólico. La H +-ATPasa de la membrana plasmática es estimulada por cationes monovalentes en el orden K+ > Na+ y es relativamente insensible a aniones. Un ejemplo de la estimulación por el K+ se ha dado en la Fig. 2.6. El calcio en general y la concentración de Ca2+ libre citosólico en particular juega un rol clave en la estimulación de esta enzima. Un incremento en el Ca2+ libre citosólico incrementa
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Los cationes son transportados “cuesta abajo” a lo largo del gradiente del potencial eléctrico (alrededor de -120 a -180mV) a través de la membrana plasmática hacia el citoplasma pon un uniporte, mediado por estructuras específicas (carriers, permeasas) en la membrana (Fig. 2.9). Para el potasio (K+), sin embargo, a bajas concentraciones externas (<1 mм) opera la alta afinidad de toma del sistema contra la diferencia del potencial electroquímico imperante (e.g., actividad exterior de 10 µm K+; actividad citoplásmica de ~80 mм K+).De este modo, se requeriría un transporte energizado de K+ cualquiera como antiporte K+-protón mediado por carriers (contratransporte) ó, mas probablemente como un simporte ó cotransporte K+-protón 1:1.
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Potencial Hídrico ( w )
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Transporte a través de membranas
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Mecanismos de absorción de nutrimentos Hidroliza ATP ATP de forma directa para transportar el compuesto en cuestión
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Movimiento de iones por por el plasmalema plasmalema y el tonoplasto
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Para el potasio (K+), sin embargo, a bajas concentraciones externas (<1mм) opera la alta afinidad de toma del sistema contra la diferencia del potencial electroquímico imperante (e.g., actividad exterior de 10 µm
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Fig. 2.10 Tasa de toma de K + (v) en función de la concentración externa de KCl (○) ó K2SO4 (∆); K = 0.023 mм). (Después de Epstein, 1972) m
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Cuando el rango de concentración es bajo, la toma frecuentemente es bien descrita por la cinética de Michaelis-Menten, como se mostró en un ejemplo en la Fig. 2.10 para la toma radical de potasio en cebada. Es evidente de esta Figura que la isoterma de la toma de potasio es la misma si la fuente de potasio es KCl o K2SO4. Como veremos después, sin embargo, cuando las concentraciones del sustrato son mayores, el anión acompañante tiene efecto en la tasa de toma del catión y viceversa. El valor Km refleja la afinidad de los centros del carrier por el ión, justo como en reacciones enzimáticas donde este indica la afinidad de la enzima por el sustrato.
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Como primera aproximación al rango de baja concentración, la cinética de Michaelis-Menten también puede emplearse para describir tasas de toma del sulfato y fosfato. Sin embargo, como resumió Jensen et al. (1987), y demostrado en unos pocos ejemplos citados abajo, es muy limitada la aplicación formal de la cinética de Michaelis-Menten por razones teóricas (Sección 2.4.2) y frecuentemente no concuerda con los resultados experimentales. El concepto original de un mecanismo de transporte iónico mediado por un solo carrier (un sistema carrier para cada ión) no describe suficientemente la cinética de la toma cuando se probaron amplios rangos de concentraciones. A concentraciones de K+ por encima de 1mм, por ejemplo, la cinética difiere considerablemente de aquella a menores concentraciones. La selectividad de los centros de ligamiento es menor (el Na+ compite con el K+) y el anión acompañante tiene un efecto sobre la tasa de toma (Sección 2.5.5). Estas diferencias conducen a la hipótesis de un sistema dual, el Sistema I con mayor
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Las desviaciones en la isoterma de la toma, cuando se considera para un amplio rango de concentración, especialmente a altas concentraciones externas fueron interpretadas como indicadores de sistemas carriers “multifásicos” o como inhibición alostérica (retroregulación negativa) de los centros del carrier a altas concentraciones celulares. En vista de usualmente muy bajas concentraciones en la solución del suelo particularmente de fósforo y potasio (Sección 13.2.2), y de los resultados de estudios en toma iónica en el rango de baja concentración (<10 μм) se introdujo el termino (Cmin) definiendo la concentración a la cual la toma iónica neta cesa
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Fig. 2.11 Presentación esquemática de las relaciones entre la tasa de toma (influjo neto = I) iónica y su concentración externa; C min = toma neta cero (influjo = eflujo).
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El eflujo de iones y otros solutos no solo es afectado por el electropotencial transmembranal y la integridad de la membrana plasmática sino también por la concentración y actividad de los respectivos
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Estos parámetros de la cinética de la toma de iones también son fuertemente afectados por el estado nutricional vegetal. Esto es cierto no solo para Cmin sino también para Km y particularmente para Imax. Se da
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Los inhibidores de la síntesis proteica deprimen fuertemente la formación de este sistema de transporte inducible en el cual los grupos
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Reducción y Asimilación del Nitrato Además de su función en la inducción de la síntesis de nitrato reductasa, el nitrato, junto con la luz, puede actuar como una “señal” que altera el particionamiento del flujo del carbono fotosintético en las hojas (Fig. 8.3). Para la asimilación del amoníaco hay una alta demanda de esqueletos de carbono y se puede asumir que se presenta competencia entre la síntesis de sacarosa y la síntesis de aminoácidos como se ha mostrado en hojas de tomate. El flujo de carbono entre ambas vías parece estar regulada vía proteínquinasas citosólicas que modulan la actividad de dos enzimas clave, la sacarosa-P sintasa y la PEP carboxilasa, mediante la fosforilación. El nitrato por si mismo funciona como una metabolito señal (Fig. 8.3). Sin embargo, ambas enzimas responden a la fosforilación en un modo inverso, la sacarosa-P sintasa es inactivada y la PEP carboxilasa activada, y por lo tanto hay un particionamiento de fotosintatos desde la síntesis preferencial de sacarosa hacía la síntesis de aminoácidos. Se requiere una alta actividad de PEP
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Reducción y Asimilación del Nitrato Fig. 8.3 Modelo del NO y luz como “señales” para la fosforilación proteica y desactivación de sacarosa-P sintasa y activación de PEP carboxilasa. (Modificado de Champigny & Foyer, 1992; reimpreso con permiso de la American Society of Plant Physiologists.)
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Reducción y Asimilación del Nitrato Localización en Raíces y Vástagos. En la mayoría de especies vegetales ambos raíces y vástagos son capaces de la reducción del nitrato, y las raíces pueden reducir entre un 5 y 95% del nitrato tomado. La proporción de la reducción llevada a cabo en las raíces y vástagos depende de varios factores, que incluyen el nivel del suministro de nitrato, de la especie vegetal, de la edad vegetal, y tiene importantes consecuencias para la nutrición mineral y la economía del carbono vegetal. En general, cuando el suministro externo de nitrato es bajo, una alta proporción del nitrato es reducida en las raíces. Con un suministro creciente de nitrato, la capacidad para la reducción del nitrato en las raíces se vuelve un factor limitante y una proporción creciente del nitrógeno total es translocada a los vástagos en la forma de nitrato (Fig. 8.4). Existen notables diferencias entre especies vegetales en ambas, la proporción de nitrato reducido en las raíces y la respuesta a crecientes concentraciones externas de nitrato. Por ejemplo, a un suministro de 1 mм nitrato la proporción del nitrógeno en la savia del xilema como nitrato fue de 15% en trébol blanco (Trifolium repens),
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Reducción y Asimilación del Nitrato Localización en Raíces y Vástagos. En la mayoría de especies vegetales ambos raíces y vástagos son capaces de la reducción del nitrato, y las raíces pueden reducir entre un 5 y 95% del nitrato tomado. La proporción de la reducción llevada a cabo en las raíces y vástagos depende de varios factores, que incluyen el nivel del suministro de nitrato, de la especie vegetal, de la edad vegetal, y tiene importantes consecuencias para la nutrición mineral y la economía del carbono vegetal. En general, cuando el suministro externo de nitrato es bajo, una alta proporción del nitrato es reducida en las raíces. Con un suministro creciente de nitrato, la capacidad para la reducción del nitrato en las raíces se vuelve un factor limitante y una proporción creciente del nitrógeno total es translocada a los vástagos en la forma de nitrato (Fig. 8.4).
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Reducción y Asimilación del Nitrato Existen notables diferencias entre especies vegetales en ambas, la proporción de nitrato reducido en las raíces y la respuesta a crecientes concentraciones externas de nitrato. Por ejemplo, a un suministro de 1 mм nitrato la proporción del nitrógeno en la savia del xilema como nitrato fue de 15% en trébol blanco (Trifolium repens), 33% en arveja (Pisum sativum) y 59% en Xanthium stumarium, y a 10 mм nitrato estas proporciones se incrementaron a 66%, 41% y 69%, respectivamente. Sin embargo, como se discutió en la Sección 3.4.4 el reciclaje del nitrógeno reducido puede ser un componente importante del nitrógeno reducido encontrado en la savia del xilema y de esta manera, el porcentaje del N total como nitrato en la savia del xilema no es un parámetro muy fiable sobre la proporción de la reducción del nitrato en la raíz. No obstante, hay un patrón general entre especies vegetales en el particionamiento de la reducción y asimilación del nitrato entre las raíces y vástagos.
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Absorción de Nitrato por las Raíces
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Absorción de Nitrato por las Raíces
Absorción del nitrato por las raíces La absorción de NO3- está sujeta a una regulación positiva o de inducción y negativa. La última parece depender del nivel de N de la planta. Ha sido sugerido que el ciclo de de los aminoácidos entre los tallos y las raíces sirve para proveer la información necesaria respecto del nivel de N en la planta, que le permite a las raíces regular la
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Transportadores •
En las plantas superiores la absorción está mediada por al menos 3 sistemas transportadores distintos que co-existen en las células radiculares.
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CHATS CHA TS (Constitutive High Affinity transport System)
•
Se caracterizan por su bajos valores de Km (6-20µM) y de Vmax (0.3-0.82 µmol g. h-1)
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IHATS (Inducible High Affinity Transport System)
• •
Km (20-100µM) y de Vmax (3-8 µmol g. h-1) se inducen dentro de las horas o días posteriores a la exposición
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LATS LA TS (Low Affinit Affinityy Transport System)
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