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La Vida En Río Tinto - Ángeles Aguilera / Virginia Souza-Egipsy / Elena González-Toril COLECCIÓN CIENCINTA

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La Vida En Río Tinto - Ángeles Aguilera / Virginia Souza-Egipsy / Elena González-Toril COLECCIÓN CIENCINTA La vida en Río Tinto - Ángeles Aguilera / Virginia Souza-Egipsy / Elena González-Toril COLECCIÓN CIENCINTA Nº1 -2020 Ángeles Aguilera / Virginia Souza-Egipsy / Elena González-Toril La vida enRí0 Tinto COLECCIÓN CIENCINTA Nº1-2020 Editorial La colección CIENCINTA pretende difundir y divul- gar los interesantes resultados de investigación del INTA, de la mano de prestigiosos investigadores, que han logrado ser referentes en sus respectivos campos de investigación. El carácter multidisciplinar del INTA ofrece la posi- bilidad de una colección divulgativa, rica, variada y atractiva, que hará las delicias de todo aquel lector que se acerque a esta apasionante aventura del saber: podrá viajar al espacio exterior, pasando por desiertos inhóspitos, descubriendo virus emergen- tes, y vida en condiciones extremas. ¡Bienvenido a Ciencinta! Miguel Ángel Alonso Editor CAB La vida en Río Tinto Ángeles Aguilera1*, Virginia Souza-Egipsy2 y Elena González-Toril1 Centro de Astrobiología-CSIC-INTA, Carretera Ajalvir Km4, 28850 Torrejón de Ardoz, España. 1 Instituto Estructura de la Materia CSIC, Serrano 113 bis, 28006, Madrid. *Corresponding Author: A. Aguilera ([email protected]), Teléfono: +34 91 5206434. CATÁLOGO GENERAL DE PUBLICACIONES OFICIALES https://cpage.mpr.gob.es Edita: © Autor y editor, 2020 NIPO: 078-20-003-2 (edición en papel) NIPO: 078-20-004-8 (edición en línea) ISBN: 978-84-9091-517-2 (edición en papel) Depósito Legal: M-21173-2020 Fecha de edición: noviembre 2020 Maquetación e impresión: Vicente Aparisi - Din Impresores ©Portada: Luis Cuesta / Laguna Ácida de Peña del Hierro Las opiniones emitidas en esta publicación son exclusiva responsabilidad del autor de la misma. Los derechos de explotación de esta obra están amparados por la Ley de Propiedad Intelectual. Ninguna de las partes de la misma puede ser reproducida, almacenada ni transmitida en ninguna forma ni por medio alguno, electrónico, mecánico o de grabación, incluido fotocopias, o por cualquier otra forma, sin permiso previo, expreso y por escrito de los titulares del © Copyright. En esta edición se ha utilizado papel 100% libre de cloro procedente de bosques gestionados de forma sostenible. Prefacio Los ecosistemas extremos y los microorganismos que los habitan, denomina- dos extremófilos, han atraído desde siempre un interés científico considerable, no sólo porque demuestran que la vida es posible y adaptable a condiciones am- bientales muy diversas y adversas sino también porque el estudio de este tipo de ambientes es imprescindible en el campo de la Astrobiología. Estos hábitats nos permiten entender mejor cuáles son los límites de la vida tal y como la co- nocemos, sirviendo como banco de pruebas a la hora de buscar vida en otras partes del universo. En este sentido, el estudio de los ambientes extremos ácidos (pH<3) es cada vez más importante ya que los ambientes ácidos son considerados como análogos terrestres de Marte, planeta al que se encaminan gran parte de las misiones espaciales actuales. Los extremófilos viven en hábitats que son muy hostiles o incluso letales para otras formas de vida y su estudio ha proporciona- do descubrimientos innovadores que desafían algunos de los paradigmas de la biología moderna. De entre los ecosistemas extremos, los ambientes ácidos son relativamente escasos en todo el mundo y están generalmente asociados con la actividad volcánica y con depósitos minerales de gran interés económico. Además de la alta acidez, estos ambientes presentan condiciones químicas y físicas muy variables, dependiendo de su origen y de la naturaleza geoquímica del lugar. El pH ácido facilita la disolución de metales pesados en el agua, lo que provoca un impacto ambiental enorme. Río Tinto, situado en el suroeste de España, es uno de los ecosistemas ácidos extremos más inusuales descritos hasta el momento debido principalmente, además de a su acidez, a su extenso tamaño (92 Km), a sus altas concentraciones de metales pesados disueltos y, sobre todo, su alto nivel de diversidad microbiana, principalmente eucariótica. Numerosos estudios sobre Río Tinto han demostrado la importancia del ciclo del hierro, no sólo en la generación de las condiciones extremas del hábitat sino también en el mantenimiento de su diversidad microbiana. A lo largo del libro, revisaremos los trabajos realizados en cuanto a la geomicrobiología, biodiversidad y ecofisiología de los microorga- nismos que habitan en uno de los mayores y más importantes ambientes ácidos extremos descritos hasta ahora. Ángeles Aguilera Elena González-Toril Virginia Souza-Egipsy Contenidos 1.- Introducción …………………………………………………………………………………………………… 7 1.1 Ambientes extremos y extremófilo ……………………………………………………………………………7 1.2 Microorganismos procariotas en ambientes extremos ……………………………………………8 1.3 Microorganismos eucariotas en ambientes extremos ………………………………………… 10 1.4 Mecanismos de resistencia en extremófilo …………………………………………………………… 13 1.5 Interés biotecnológico de los extremófilo …………………………………………………………… 16 2. Naturaleza y origen de los ecosistemas ácidos extremos ……………………19 3. Río Tinto y su entorno ……………………………………………………………………………………27 3.1 Contexto geológico: Faja Pirítica Ibérica ………………………………………………………………… 27 3.2 Cuenca hidrológica y clima ……………………………………………………………………………………… 28 3.3 Comparación con otros ambientes ácidos extremos …………………………………………… 32 4.- Biodiversidad microbiana de Río Tinto ……………………………………………………35 4.1 Biodiversidad procariota …………………………………………………………………………………………… 36 4.2 Biodiversidad eucariota …………………………………………………………………………………………… 43 4.2.1 Microalgas acidófila ………………………………………………………………………………………46 4.2.2 Euglenófitos y protozoos acidófil ………………………………………………………………48 4.2.3 Hongos y levaduras acidófila …………………………………………………………………………48 4.2.4 Plantas ………………………………………………………………………………………………………………49 4.3 Dinámica poblacional de las comunidades eucariotas ………………………………………… 51 5.- Modelo geomicrobiológico de Río Tinto …………………………………………………53 5.1 Modelo geomicrobiológico del cauce principal ……………………………………………………… 53 5.2 Modelo geomicrobiológico de las salidas de mina ……………………………………………… 56 5.3 Modelo geomicrobiológico de los lagos de cortas de mina ………………………………… 57 5.4 Papel de los eucariotas en el modelo geomicrobiológico de aguas ácidas ………… 61 Contenidos 6.- Formación y estructura de los biofilm ………………………………………………… 63 7.- Mecanismos de adaptación a ambientes ácidos extremos ………………… 71 7.1 Mecanismos de adaptación de procariotas acidófilo …………………………………………… 71 7.2 Mecanismos de adaptación de eucariotas acidófilo …………………………………………… 75 7.2.1 Adaptaciones a pH ácidos ……………………………………………………………………………… 75 7.2.2 Adaptaciones a limitaciones nutricionales ………………………………………………… 76 7.2.3 Adaptaciones fotosintéticas ………………………………………………………………………… 77 7.2.4 Adaptaciones a metales pesados …………………………………………………………………… 81 7.3 La transferencia horizontal de genes como mecanismo de resistencia adaptativa …………………………………………………………………………………………84 8.- Río Tinto y su importancia en Astrobiología ……………………………………… 87 8.1 Interés astrobiológico de los extremófilos y los ambientes extremo …………………89 8.2 Río Tinto como análogo terrestre de Marte ……………………………………………………………90 9.- Bibliografía ………………………………………………………………………………………………… 93 1. Introducción 1.1 Ambientes extremos consideramos normal y los organis- y extremófilos mos que viven en los límites de esta normalidad ambiental o más allá son La exploración del planeta nos ha lle- considerados extremófilos. Es intere- vado a continuos descubrimientos de sante señalar que los extremófilos no vida en ambientes que, inicialmente solo soportan tales condiciones ad- fueron considerados como inhabita- versas para llevar a cabo su actividad bles. De hecho, la vida puede surgir y metabólica, si no que necesitan de prosperar bajo condiciones ambienta- dichas condiciones extremas para de- 7 les muy duras. Esto ha dado lugar al sarrollarse. Los microorganismos capa- origen de una nueva disciplina dentro ces de tolerar estas condiciones, pero de la biología, la Extremofília, que es- que también pueden desarrollarse en tudia la diversidad, ecología y fisiol - hábitats moderados, no son verdade- gía de los ambientes extremos y los ros extremófilos, siendo considerados organismos que los habitan. Estos eco- extremo-tolerantes. sistemas se definen como lugares que El número de extremófilos iden- tienen uno o varios factores ambien- tificados ha crecido rápidamente en tales incompatibles con la mayoría de los últimos años (Tabla 1). Algunos las formas de vida, especialmente con ejemplos de esto los podemos encon- la vida humana, lo que la convierte trar en las comunidades microbianas en una defi ición absolutamente an- que viven a temperaturas elevadas, tropocéntrica. A pesar de la aparente como es el caso de las arqueas hiper- hostilidad de dichos hábitats, los eco- termófilas que pueden encontrarse a sistemas extremos pueden contener temperaturas superiores a los 100ºC un nivel de biodiversidad mayor de [1]. Otros organismos viven en zonas lo esperado. El medio ambiente que polares frías, como los psicrófilos de conocemos tiene oxígeno, nunca es la Antártida que son capaces de vivir demasiado frío ni demasiado caliente en aguas salobres con una salinidad (la mayoría de los organismos viven varias veces mayor que la del agua en un rango relativamente limitado de mar, a temperaturas inferiores a de temperaturas de 5°C a 40°C) una -10ºC y bajo una capa de hielo de 20 radiación moderada por la atmósfera. m de espesor [2]. De manera similar, Este mundo familiar define lo que hay organismos que viven en ambien- consisten en formas
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