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Especies Oportunistas De Importancia Clínica De Los Géneros Bipolaris

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Especies Oportunistas De Importancia Clínica De Los Géneros Bipolaris Bol. Micol. 2015; 30(2): 40-63 TAXONOMÍA Especies oportunistas de importancia clínica de los géneros Bipolaris Shoemaker y Curvularia Boedijn: su caracterización bajo los nuevos criterios taxonómicos (Opportunistic species of clinical relevance in the Bipolaris Shoemaker & Curvularia Boedijn genera: its characterization under newest taxonomic criteria) Eduardo Piontelli Laforet. Excátedra de Micología, Escuela de Medicina Universidad de Valparaíso, Casilla 92 V, Valparaíso, Chile. [email protected] RECIBIDO:16 de Noviembre de 2015 APROBADO:11 de Diciembre de 2015 EL AUTOR DECLARA NO TENER CONFLICTO DE INTERESES Palabras clave: Curvularia, Bipolaris, taxonomía Key words: Curvularia, Bipolaris, taxonomy RESUMEN el acceso a la información referente en la litera- tura moderna, que gracias a la biología molecular Los géneros fúngicos Bipolaris y Curvu- ha resuelto muchas de las dudas de la taxonomía laria, incluyen numerosas especies fitopatógenas, basada solo en el fenotipo. saprotrofas y algunas oportunistas emergentes en Las especies de ambos géneros (princi- el ser humano como en otros animales. La distribu- palmente Curvularia) pueden afectar a pacientes ción ecológica de ambos taxas es cosmopolita en inmunocomprometidos e inmunocompetentes, la gran mayoría de sus integrantes, encontrándose en especial B. cynodontis, C. australiensis C. frecuentemente en hojas, tallos y raíces de dife- hawaiensis y C. spicifera, causando cuadros clíni- rentes pastos, en especial Poaceae, pero también cos diversos ya sea superficiales o profundos en en una gran variedad de dicotiledoneas, aire y sue- muchas áreas geográficas. lo en todos los continentes. Cochliobolus, Bipola- Los objetivos de este trabajo solo preten- ris y Curvularia integran un complejo de especies den aportar para el micólogo médico dedicado al taxonómicamente confuso, debido a los constantes laboratorio, los avances más significativos en la cambios en la nomenclatura de algunos de sus in- nomenclatura, taxonomía y biología molecular de tegrantes asexuales (Bipolaris y Curvularia), los las especies comunes de importancia clínica de cuales se han diferenciado principalmente en base ambos géneros. Se incluyen además como ob- a la morfología de sus conidios, situación a veces jetivo primario, descripciones, figuras, claves y la muy dificultosa debido a que en ambos géneros literatura actual de utilidad en la identificación de algunas especies presentan similares característi- los agentes causales, sin olvidar la valoración de cas conidiales. Una cuidadosa identificación y una sus sensibilidades frente a los antifúngicos moder- nominación precisa de las especies es crucial para nos. Se aconseja además, que si el laboratorio es micologia.uv.cl 41 Especies oportunistas de importancia clínica de los géneros Bipolaris Shoemaker y Curvularia Boedijn - E. Piontelli. de baja complejidad, debe derivar su aislado a un It is also advisable, if the laboratory is low centro de referencia internacional para confirmar in complexity, must derive its isolated tu a interna- mediante biología molecular su diagnóstico. tional reference center to confirm the diagnostic by molecular biology. ABSTRACT INTRODUCCIÓN The fungal genera Bipolaris and Curvu- laria, include numerous plant pathogenic species, Shoemaker (1959), estableció el género Bi- saprophitic and some emerging opportunistic in polaris para las especies que Nisikado (1928) co- man and other animals. The ecological distribution locó en su subgenero Euhelminthosporium. Los of both taxa is cosmopolitan in the vast majority of conidios se describieron como fusoides, derechos o its members, often being in leaves, stems and roots curvos y capaces de germinar en ambos extremos of different grasses, especially Poaceae, but also in del conidio. Especie tipo: Bipolaris maydis (Nisik. a variety of dicotyledonous, air and soil on all con- & Miyake) Shoem. Teleomorfo (= Cochliobolus). tinents. Cochliobolus, Bipolaris and Curvularia Las especies de Bipolaris Shoemaker y Curvularia integrate a complex of species taxonomically con- Boedijn (Ascomycota, Dothideomycetes, Pleospo- fusing due to the constant changes in the nomen- rales, Pleosporaceae), son importantes patógenos clature of some of its asexual members (Bipolaris en más de 60 géneros de plantas hospedadoras o and Curvularia), which are differentiated mainly saprotrofas (Sivanesan 1987; Rong, 2001;Schoch based on the morphology of its conidia, a situation et al, 2009; Manamgoda et al., 2011,2012, 2014, sometimes very difficult because in both genera 2015; da Cunha et al, 2012 ; Tan et al, 2014; Ma- conidial of some species have similar characteris- drid et al, 2014), sin embargo, unas pocas especies tics. Careful identification and a precise nomina- de estos géneros son ocasionalmente causantes de tion of species is crucial for access to information infecciones oportunistas en humanos u otros ani- concerning in modern literature, actually thanks to males (Hoog et al. 2000-2011). Los frecuentes molecular biology has solved many of the doubts cambios en la nomenclatura, como resultado en los of taxonomy based only on the phenotype. avances taxonómicos, son importantes para los mi- The species of both genera (mainly Cur- cólogos de las diferentes ramas científicas, ya sea vularia) can affect immunocompetent or debilitate fitopatólogos o dedicados a las diferentes ciencias patients, especially B. cynodontis, C. australien- biológicas (Sivanesan 1987-1992; Manamgoda et sis, C. hawaiensis and C.spicifera, causing va- al. 2011, 2012, 2014). Bipolaris y Curvularia pre- rious superficial or systemic clinical cases in many sentan similares morfologías (Sivanesan 1987) y geographic areas. algunas especies de Bipolaris con conidios cortos The objectives of this paper are intended y derechos presentan características intermedias only to provide for medical mycologist dedicated entre éstos 2 géneros, a diferencia de las especies to the laboratory, the most significant developments con grandes y largos conidios curvos (Bipolaris in the nomenclature, taxonomy and molecular bio- maydis). logy of the common species of clinical importance Los estudios filogenéticos moleculares of both genera. Also included as a primary objec- recientes en Bipolaris y Curvularia, demostra- tive, descriptions, figures, keys and the current li- ron que la primera no es monofilética y algunos terature useful in identifying the causative agents, de sus integrantes con conidios cortos y derechos without forgetting the value of its senbility against se agrupan con Curvularia (Goh 1998; Berbee et modern antifungals. al. 1999; Emami & Hack ,2002; Manamgoda et Bol. Micol. 2015; 30(2): 40- 63 micologia.uv.cl 42 Especies oportunistas de importancia clínica de los géneros Bipolaris Shoemaker y Curvularia Boedijn - E. Piontelli. al. 2011-2012). El teleomorfo Pseudocochliobo- genérico se divide en 2 grupos. Las especies de lus fue primariamente separado de Cochliobolus Bipolaris y Cochliobolus se agrupan en el Grupo 1 basándose en su morfología sexual (Tsuda et al. junto a sus tipos especies, mientras las especies de 1977), pero se ha tratado generalmente como sinó- Curvularia (incluyendo especies llamadas como nimo (Alcorn,1983; Sivanesan,1987). Debido a la Bipolaris, Cochliobolus y Curvularia), se agrupan alta variación intraespecífica de algunos caracteres en el Grupo 2 con sus tipos genéricos, resolviéndo- encontrados, Alcorn (1983), dudó en su uso para se el conflicto en la nomenclatura y transfiriendo una delineación genética debido a que ambos gé- algunas especies de Bipolaris a Curvularia (19 neros producen estados asexuales en Bipolaris y Bipolaris, 25 Cochliobolus y 12 Curvularia). Sin Curvularia, lo que permitió las bases para la sino- embargo, existen 95 especies de Bipolaris, 30 de nimía entre Pseudocochliobolus y Cochliobolus. Cochliobolus y 101 de Curvularia que no han sido Ecología. Las especies de Bipolaris y Curvularia secuenciadas y por ende no adscritas a sus nuevas se aíslan con frecuencia en hojas, tallos y raíces posibles especies, para clarificar su posición. Ta- de diferentes pastos, algunas especies de Bipola- zick &, Tajick (2013), al emplear datos de análi- ris y Curvularia (B. cynodontis y C. lunata) se sis de multigenes, obtuvieron similares resultados encuentran en pasto bermuda y otros pastos o di- y confirmaron la correcta posición evolutiva y el versos vegetales Chloris, Cymbopogon, Cynodon, estatus taxonómico de Bipolaris oryzae. Manag- Hordeum, Oryza, Pennisetum. Saccharum, Sor- moda et al. (2014), agregan C.spicifera (=B. spici- gum, Zea, etc. pero también desde una gran varie- fera), sin embargo, secuencias adicionales como la dad de dicotiledoneas, aire y suelo en todos los calmodulina (cmd), el factor de elongación y trans- continentes (Sivanesan, 1987;Piontelli & Grizzoli lación 1α (tef), tubulina (tub) y los espacios inter- 1994; Picco et al, 1999). génicos del DNAr (IGS), pueden emplearse en el Biología Molecular. Un árbol genético empleando análisis. Por lo tanto, las secuencias de ITS + tef + el gen Brn1, mostró que Bipolaris se separa en 2 tub+ RAPD + IGS se combinan en una matriz de mayores clados con respecto a Pseudocochliobolus datos, los cuales se analizan empleando el método y Cochliobolus (Shimizu et al. 1998), mientras las MrBayes (Huenselbeck & Ronquist, 2001; Gomez cepas de Curvularia, se agrupan solo en el grupo 2 (Berbee et al. 1999). Un análisis filogenético en las Pleosporaceae por Kodsueb et al. (2006), revela- ron que Cochliobolus y sus anamorfos se dividen ampliamente
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