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(LAMIACEAE) DE ARGENTINA Morpho-Anatomical Study of Gynoecium and Fruit in Mentha (Lamiaceae) Species from Argentina

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(LAMIACEAE) DE ARGENTINA Morpho-Anatomical Study of Gynoecium and Fruit in Mentha (Lamiaceae) Species from Argentina BOTÁNICA-MORFOLOGÍA http://www.icn.unal.edu.co/ Caldasia 33(2):349-366.Bonzani et2011 al. ESTUDIOS MORFO-ANATÓMICOS DE GINECEO Y FRUTO EN ESPECIES DE MENTHA (LAMIACEAE) DE ARGENTINA Morpho-anatomical study of gynoecium and fruit in Mentha (Lamiaceae) species from Argentina NORMA E. BONZANI Instituto Multidisciplinario de Biología Vegetal y Facultad de Ciencias Químicas (UNC). ����C.C. 495. 5000. Córdoba, Argentina. [email protected] VIVIANA S. BRAVI Departamento de Farmacia. Facultad de Ciencias Químicas. Universidad Nacional de Córdoba (UNC), Córdoba, Argentina. [email protected] GLORIA E. BARBOZA Instituto Multidisciplinario de Biología Vegetal y Facultad de Ciencias Químicas (UNC). C.C. 495. 5000. Córdoba, Argentina. [email protected] RESUMEN Se realizaron estudios morfológicos y anatómicos de gineceo y fruto, utilizando microscopía óptica y electrónica de barrido, en especies de Mentha (sect. Mentha L.y sect. Pulegium (Mill.) Coss. & Germ.). Se registran por primera vez para el género algunas novedades sobre el gineceo, histología del nectario, microesculturas en el epicarpo, presencia de ceras epicuticulares en el hilo y la morfología de los cristaloides. Se establecieron cuatro tipos de microesculturas en núculas maduras. Los resultados son evaluados desde el punto de vista taxonómico para la delimitación de las especies. Además, se registra por primera vez mixocarpia en M. citrata, M. x piperita y M. spicata y se plantean algunas consideraciones sobre este fenómeno. Palabras clave. Nectario, núculas, mixocarpia, ceras epicuticulares, Mentheae. ABSTRACT Morphological and anatomical studies on the gynoecium and fruit of Mentha species (sect. Mentha L. and sect. Pulegium (Mill.) Coss. & Germ.) from Argentina were done using optic and scanning electronic microscope. Some novelties on the gynoecium, nectary histology, microsculpture of the nutlets, epicuticular waxes in the hilum and the morphology of the crystalloids are reported for the first time for Mentha. Four types of microsculpture on the surface of the nutlets were established. The results are evaluated from a taxonomical point of view for species delimitation. In addition, myxocarpy is reported for M. citrata, M. x piperita and M. spicata for the first time and its occurrence is discussed. Key words. Nectary, nutlets, myxocarpy, epicuticular waxes, Mentheae. 349 Morfoanatomía de gineceo y fruto en Mentha INTRODUCCIÓN Desde el punto de vista exomorfológico, la presencia de ceras epicuticulares depositadas El género Mentha L. (fam. Lamiaceae, subfam. en el hilo sólo son mencionadas para otros Nepetoideae, tribu Mentheae), originario de taxones de Lamiaceae (Martín-Mosquero Europa y Asia, se halla naturalizado en et al. 2004), mientras que el estudio de las América encontrándose en Argentina tres microesculturas de la superficie de las núculas especies y dos híbridos (Bonzani et al. 2007). ha recibido más atención por su relevancia Estas especies pertenecientes a las sect. taxonómica (Salmaki et al. 2008, Marin et Mentha (Mentha citrata Ehrh., M. spicata al. 1994, Martín-Mosquero et al. 2006a,b, L., M. x piperita L. y M. x rotundifolia (L.) Moon & Hong 2006). Por último, Bouman Hudson y a la sect. Pulegium: M. pulegium & Meeuse (1992) destacan la importancia de L., son muy apreciadas por sus propiedades estos caracteres como estrategia de dispersión terapéuticas y por su alto contenido en aceites y hacen referencia, entre otros taxones, a esenciales, se utilizan en preparaciones Mentha aquatica L. farmacéuticas y como fitoterápicos, además de ser expendidos en el comercio como La mixocarpia, que consiste en la liberación medicamento herbolario. Por sus propiedades de mucílago por las células del epicarpo de aromáticas y saporíferas su uso se extiende las núculas maduras en contacto con el agua, también a la repostería, licorería e industria es otro aspecto interesante de Lamiaceae de las golosinas (Bruneton 1991, Alonso reportada en varios géneros, incluido Mentha 1998, 2004, Lawrence 2006). (Witzthum 1978, Ryding 1992a, 2001, Duletiæ-Lauševiæ & Marin 1999, Fernández- De los estudios taxonómicos del género Alonso et al. 2009). se destacan los trabajos de Harley (1972), Harley & Brighton (1977), Mabberley (1997) A fin de continuar contribuyendo con el y Gober et al. (2002), reconociéndose hasta conocimiento del género, abordamos en esta el momento 25 especies y varios híbridos oportunidad el estudio de las estructuras (Harley & Brighton 1977). Estudios previos reproductivas con los siguientes objetivos: 1) se han referido a la morfo-anatomía de conocer la estructura del gineceo abarcando los órganos aéreos (tallos, hojas) y a las un detallado estudio del nectario, 2) proveer estructuras secretoras de Mentha, fuentes una descripción de la morfo-anatomía de las de obtención de aceites esenciales (Clauss núculas, incluyendo las microesculturas de su & Tyler 1968, Muñoz Centeno et al. 1998, superficie, 3) describir la morfología de las Martins 2002, Martins & Martins 2003, ceras epicuticulares depositadas en el hilo, Bonzani et al. 2007). Por su parte, las 4) comprobar la presencia de mixocarpia en estructuras reproductivas, es decir ovario, núculas maduras, 5) evaluar el significado nectario, fruto y órganos de dispersión taxonómico de las variaciones observadas en (núculas) fueron descritas por Hedge (1970) tales caracteres. y Cronquist (1981) para Lamiaceae en general. MATERIALES Y MÉTODOS Con respecto a la estructura anatómica del Se trabajó con materiales frescos y pericarpo se destacan los trabajos de Moon herborizados, todos provenientes de Argentina & Hong (2006), Duletiae-Lauševiaæ & Marin y depositados en el Herbario del Museo (1999) y Ryding (1992a,b, 1993, 1995, 2008) Botánico de Córdoba (CORD), Argentina. Las quienes, junto a otros géneros de Lamiaceae, especies analizadas fueron: M. citrata Ehrh.: analizan algunas especies de Mentha. Prov. Córdoba, Dpto. Colón, Salsipuedes, 29 350 Bonzani et al. set 1996, Ariza Espinar 3420 y 3421; Prov. se realizó con hipoclorito de sodio al 30 % Córdoba, Dpto. Capital, Bower, 30 mar 2008, y posterior lavado con agua destilada. Para Bonzani 118. seguir el recorrido de los haces vasculares se M. spicata L.: Prov. Córdoba, Dpto. Colón, tiñó con safranina acuosa al 1%. El contenido Salsipuedes, 17 ene 2000, Bonzani 83; de almidón del nectario se evidenció con Prov. Córdoba, Dpto. Capital, Barrio Urca, solución de lugol (iodo-ioduro de potasio). 15 feb 2000, Bonzani 88; Prov. San Juan, Para describir la organización histológica del Dpto. Caucete, 21 dic 1980, Hunziker & nectario se siguió la terminología propuesta Subils 23956. por Nepi (2007). M. pulegium L.: Prov. Córdoba, Dpto. Colón, Saldán, San Remo, 21 ene 2000, Bonzani 86; Se empleó también el microscopio electrónico Prov. Buenos Aires, Partido J. Madariaga, de barrido (MEB) para el análisis del gineceo Ruta 74 entre Gral. Madariaga y Las Armas, y superficie de núculas maduras. Se aplicó 21 ene 1973, Di Fulvio 309. la técnica de Gesterberger & Leins (1978), M. x piperita L.: Prov. Córdoba, Dpto. Colón, secado a punto crítico, montaje sobre discos Saldán, Los Alazanes, 21 ene 2000, Bonzani de aluminio con cinta adhesiva doble faz y 84 y Bonzani 88; Prov. San Juan, Dpto. posterior baño de oro. Chimbas, 6 ene 1961, Ariza Espinar 1298. M. x rotundifolia (L.) Hudson: Prov. Córdoba, Para evidenciar la producción de mucílago Dpto. Colón, Salsipuedes, 24 ene 2000, en las núculas maduras se siguieron dos Bonzani 87; Prov. Córdoba, Dpto. Capital, procedimientos: a) se prepararon cámaras Bower, 2 dic 2002, Bonzani 93; Prov. San húmedas montadas en placas de Petri, con Juan, Dpto. Chimbas, 7 ene 1960, Ariza papel de filtro embebido en agua destilada y Espinar 666. mantenidas con luz a una temperatura de 25- 27° C, siguiendo la técnica de Alburquerque Se realizaron preparados histológicos & Andrade (1998); b) se sumergieron las permanentes; el material fresco (ovario y núculas directamente en agua destilada en núculas) fue fijado en FAA (formol, alcohol tubos Eppendorf de 1,5 ml por un tiempo etílico y ácido acético, 3:1:1) al que se le máximo de 72 h. En ambos casos el mucílago aplicaron luego las técnicas de deshidratación fue puesto en evidencia con soluciones e inclusión en parafina para la obtención acuosas de azul de metileno al 0,05% y azul de cortes microtómicos de 10 a 14 µm. Las brillante de cresilo al 1%. El ancho del halo tinciones fueron realizadas con azul brillante mucilaginoso se determinó por mediciones de cresilo, en solución acuosa al 0,05 % (Pérez directas, expresadas en milímetros. Para et al. 2002), safranina alcohólica al 1% y fast- tipificar la reacción de mixocarpia en las green alcohólico al 0,1% (Martín-Mosquero especies estudiadas se tomaron los parámetros et al. 2004). Posteriormente, se procedió al propuestos por Duletiæ-Lauševiæ & Marin montaje con bálsamo de Canadá (Johansen (1999), a saber: reacción fuerte (mayor de 1940, D’Ambrogio de Argüeso 1986). Para 0,5 mm), moderadamente fuerte (entre 0,1- el estudio del pericarpo las núculas maduras 0,5 mm), débil (menor a 0,1 mm) y muy fueron sumergidas 24 h en agua destilada y débil (menor a 0,05 mm). Para definir las glicerol (90:10) antes de la aplicación de las microesculturas de las núculas se tomaron en técnicas clásicas histológicas. consideración los modelos de Barthlott (1981) y Barthlott et al. (1998). Para el análisis del receptáculo se efectuó la diafanización del mismo desprovisto de Las ilustraciones se realizaron con antófilos estériles y estambres. El diafanizado fotomicroscopio
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