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PROTISTAS MARINOS Viviana A PROTISTAS MARINOS Viviana A. Alder INTRODUCCIÓN plantas y animales. Según este esquema básico, a las plantas les correspondían las características de En 1673, el editor de Philosophical Transac- ser organismos sésiles con pigmentos fotosinté- tions of the Royal Society of London recibió una ticos para la síntesis de las sustancias esenciales carta del anatomista Regnier de Graaf informan- para su metabolismo a partir de sustancias inor- do que un comerciante holandés, Antonie van gánicas (nutrición autótrofa), y de poseer células Leeuwenhoek, había “diseñado microscopios rodeadas por paredes de celulosa. En oposición muy superiores a aquéllos que hemos visto has- a las plantas, les correspondía a los animales los ta ahora”. Van Leeuwenhoek vendía lana, algo- atributos de tener motilidad activa y de carecer dón y otros materiales textiles, y se había visto tanto de pigmentos fotosintéticos (debiendo por en la necesidad de mejorar las lentes de aumento lo tanto procurarse su alimento a partir de sustan- que comúnmente usaba para contar el número cias orgánicas sintetizadas por otros organismos) de hebras y evaluar la calidad de fibras y tejidos. como de paredes celulósicas en sus células. Así fue que construyó su primer microscopio de Es a partir de los estudios de Georg Gol- lente única: simple, pequeño, pero con un poder dfuss (1782-1848) que estos diminutos organis- de magnificación de hasta 300 aumentos (¡diez mos, invisibles a ojo desnudo, comienzan a ser veces más que sus precursores!). Este magnífico clasificados como plantas primarias (protofitas) dispositivo le permitió no sólo lograr una ópti- o animales primarios (protozoos). Este nuevo sis- ma eficiencia en su trabajo sino que, al utilizarlo tema de clasificación permaneció en uso e inal- para observar el agua de una laguna, descubrió un terable durante muchos años, a pesar de que las mundo insospechado y fascinante: una multitud formas de vida que iban descubriéndose no se de “animálculos” y “pequeñas anguilas” entrecru- ajustaban a las características empleadas para di- zándose y nadando en todas direcciones. En sus ferenciar entre plantas y animales, por lo que su cartas a la Royal Society figuran pues las prime- asignación a uno u otro reino resultaba forzada. ras descripciones de lo que hoy conocemos como En 1866, el zoólogo alemán Ernst Haeckel pro- bacterias y protozoos, abriendo así el camino a la pone la creación de un tercer reino, Protista, para microbiología y a la protistología. Más tarde, van incluir a todos aquellos organismos “inferiores” Leeuwenhoek complementaría sus estudios con que no satisfacían los requerimientos para ser observaciones sobre espermatozoides, células incorporados a ninguno de los otros dos reinos. sanguíneas y diversos temas de embriología, his- Este concepto, desatendido por la gran mayoría tología, entomología, botánica y cristalografía. Su de los investigadores de la época, originalmente arte para la construcción de microscopios residía abarcaba no sólo a las algas y los protozoos sino en la excelente calidad del cristal empleado y en también a las bacterias. Posteriormente, Copeland la técnica de pulido del mismo para lograr lentes (1938, 1956) propone ubicar a los microorganis- biconvexas de corta distancia focal. Esta técnica mos sin núcleo (bacterias y algas azules) en un fue mantenida en celoso secreto por van Leeuw- reino aparte al que llama Monera, y unos años enhoek hasta su muerte, motivo por el cual las si- más tarde Whittaker (1969), apoyándose en sus guientes observaciones de microorganismos que propias observaciones ecológicas y en los trabajos lograron realizarse fueron recién a comienzos del de Stanier y van Niel (1962) sobre la diferencia- siglo XIX, gracias al advenimiento del microsco- ción entre organismos eucariotas y procariotas, pio compuesto. Desde la antigüedad hasta enton- crea un sistema de cinco reinos: Monera, Protista, ces, el universo biótico se dividía simplemente en Fungi, Plantae y Animalia. 13 Protistas Marinos Posteriormente, el uso de la microscopía elec- flagelados heterótrofos, incluidos los Euglenozoa trónica permitió establecer nuevas relaciones y Heterolobosea; filogenéticas entre varios taxones mayores de (3) SAR, que reúne a Stramenopiles (algas protistas mediante el reconocimiento de organe- pardas, diatomeas, crisofíceas, xantofíceas), Al- las presumiblemente homólogas. Asimismo, las veolata (ciliados, dinoflagelados, Apicomplexa) características bioquímicas y fisiológicas de los y Rhizaria (foraminíferos, radiolarios, cercozoos organismos, a veces vinculadas a las organelas con filopodios); cuya ultraestructura se estaba revelando, también (4) OPISTHOKONTA, que agrupa a los me- contribuyeron a echar luz sobre las afinidades tazoos (animales con tejidos diferenciados), los propuestas. A esta altura de los acontecimientos, hongos, los coanoflagelados y los Mesomyceto- sin embargo, varios grupos de protistas ya ha- zoa; y bían sido incluidos simultáneamente tanto en el (5) ARCHAEPLASTIDA, que agrupa a las Código Internacional de Nomenclatura Botánica Glaucophyta, las algas rojas, las algas verdes y las (ICBN) como en el Código Internacional de No- plantas superiores. menclatura Zoológica (IBZN), registrándose la La definición actual de “protistas” los caracte- descripción paralela de los mismos taxones con riza como organismos eucariotas de organización nombres distintos. Durante las décadas de 1970 unicelular, colonial, filamentosa o parenquimato- y 1980 la situación se tornó aún más confusa por sa, pero en todos los casos sin tejidos vegetativos la reclasificación de muchos géneros sobre la diferenciados, salvo para estructuras intervinien- base de nuevas técnicas. Esto generó cambios en tes en la reproducción (Adl et al., 2005). Esta de- los sistemas de clasificación tradicionales de los finición no responde a un esquema taxonómico, eucariotas en general, aunque los protistas fue- ya que podemos encontrar organismos con estas ron los más afectados en virtud del permanente características en los cinco conjuntos menciona- descubrimiento de nuevos taxones en base a es- dos anteriormente. En la actualidad, la idea de un tudios filogenéticos moleculares. Estos estudios, reino independiente y exclusivo para los protistas, actualmente en gran auge, implican el análisis de si bien atractiva, fue descartada por la gran mayo- ciertas secuencias de ADN, principalmente las de ría de los protistólogos. Sin embargo, el término los genes que codifican para ARN ribosómico, sigue empleándose en forma corriente por resultar como es el caso de la subunidad 18S ADNr. Las práctico e ilustrativo, sobre todo desde el punto ventajas que ofrece esta subunidad de ADNr son de vista pedagógico, y se emplea a menudo para varias: está presente en todos los eucariotas con referirse exclusivamente a los eucariotas unicelu- muchas copias por genoma, permitiendo así tra- lares. bajar al nivel de individuo; posee un alto nivel de La célula típica de un protista posee las orga- expresión, lo cual permite encarar investigaciones nelas propias de una célula eucariota (núcleo, ecológicas moleculares al nivel del ARN; y com- mitocondrias, membrana plasmática, aparato de prende un mosaico de secuencias nucleotídicas Golgi, retículo endoplasmático, vacuolas contrác- que alterna entre algunas altamente conservadas tiles, plástidos, lisosomas) más otras organelas o y otras muy variables, permitiendo así la recons- estructuras vinculadas al hecho de ser organismos trucción filogenética a diferentes niveles taxonó- completos y autónomos (flagelos, cilias, pseudo- micos (Pawlowski et al., 2012). podios, citostoma, citoprocto, vacuolas alimenti- Los estudios filogenéticos moleculares han cias, manchas oculares/estigmas, estructuras es- dado origen a un esquema de clasificación jerár- queletarias, extrusomas). El núcleo de los protis- quica de los organismos donde se elude el uso de tas contiene ADN asociado a proteínas, formando rangos formales superiores (Simpson y Roger, cromosomas. La reproducción es asexual, pero en 2002; Adl et al., 2005, 2012), optándose por di- la gran mayoría de los grupos estudiados se ha vidir a todos los organismos eucariotas, incluidos verificado también actividad sexual con procesos los protistas, en cinco grandes conjuntos o super- mitóticos y meióticos. Los organismos pueden ser grupos: de vida libre o simbióticos (mutualistas, comensa- (1) AMOEBOZOA, que agrupa a las amebas les o parásitos). Su ciclo de vida a menudo implica tradicionales, los mohos mucilaginosos, varias la formación de quistes (estadios de reposo o de tecamebas y algunos ameboflagelados; resistencia) o esporas (estadios de multiplicación (2) EXCAVATA, que agrupa a varios grupos de y propagación). Pueden ser autótrofos (no trata- 14 Protistas Marinos dos en este Capítulo), heterótrofos o mixótrofos, términos, de aquí en adelante nos referiremos a configurando un grupo polifilético sin un ancestro estos organismos simplemente como protistas. común. Todos los protistas son aerobios, vale de- cir que usan oxígeno para extraer la energía de las DISTRIBUCIÓN GEOGRÁFICA sustancias orgánicas. Si bien hasta épocas recien- tes se consideraba que existían varias especies se- Los protistas marinos se encuentran amplia- cundariamente anaerobias (tras haberse adaptado, mente representados tanto en los ambientes pe- en el curso de la evolución, a ambientes pobres en lágicos como bentónicos. La distribución de las oxígeno), actualmente se cree que la anaerobiosis formas de vida libre en la columna de agua va- en estos organismos es una
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