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Rhizome and Root Anatomy of 14 Species of Bromeliaceae 1

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Rhizome and Root Anatomy of 14 Species of Bromeliaceae 1 RHIZOME AND ROOT ANATOMY OF 14 SPECIES OF BROMELIACEAE 1 Suzana Lúcia Proença2,3,4 & Maria das Graças Sajo2,3 ABSTRACT (Rhizome and root anatomy of 14 species of Bromeliaceae) The anatomy of rhizomes and roots of 14 species of Bromeliaceae that occur in the cerrado biome were studied with the aim of pointing out particular anatomical features of the family and possible adaptations related to the environment. All the rhizomes are similar although some have root regions growing inside the cortex. In some species the vascular cylinder of the rhizome is clearly limited from the cortex. The roots are also very similar, although the coating tissue differs in roots growing inside the rhizome or externally to it and the cortex has a variable organization according to the region. The studied species present anatomical features that are associated to water absorption and storage, showing that they are adapted to the cerrado environment. Key words: bromeliads, Pitcairnioideae, Bromelioideae, Tillandsioideae, water capture, water retention, ‘cerrado’. RESUMO (Anatomia de raízes e rizomas de 14 espécies de Bromeliaceae) Com o objetivo de reconhecer caracteres particulares de Bromeliaceae e indicar possíveis formas de adaptação ao ambiente, foi estudada a anatomia dos rizomas e raízes de 14 espécies de Bromeliaceae que ocorrem no cerrado. Os rizomas apresentam estrutura básica semelhante, embora alguns deles possuam porções radiculares crescendo no interior de seu córtex. De acordo com a espécie considerada, os rizomas podem apresentar um cilindro vascular de delimitação mais ou menos nítida. As raízes também possuem estrutura básica semelhante, apesar do tecido de revestimento variar de acordo com a porção analisada (dentro do rizoma ou externa). Além disso, as raízes apresentam uma região cortical de organização variada, de acordo com a região do órgão, e um cilindro vascular. Muitos dos caracteres anatômicos observados estão associados a mecanismos de captação e retenção hídrica, mostrando que as espécies estudadas estão adaptadas ao ambiente de cerrado. Palavras-chave: bromélias, Pitcairnioideae, Bromelioideae, Tillandsioideae, captura de água, retenção hídrica, cerrado. INTRODUCTION Bromeliaceae has traditionally been divided Bromeliaceae is included in the order in three subfamilies: Pitcairnioideae, Poales (APG II 2003) and comprises 56 Tillandsioideae and Bromelioideae (Smith & genera and around 3000 species (Luther Downs 1974, 1977, 1979; Dahlgren et al. 2002) that, except for Pitcairnia feliciana 1985), although recent phylogenetic (A. Chev.) Harms & Mildbr. from the West analyses do not confirm the monophyly of Africa, occupy various habitats in the Pitcairnioideae (Crayn et al. 2000; Horres tropical and subtropical regions of the New et al. 2000). The subfamilies are separated World (Pittendrigh 1948). according to the growth habit, the fruit and The plants are mostly herbaceous with seed morphology and the ovary position a reduced stem from where leaves, (Smith & Downs 1974, 1977, 1979). inflorescences and lateral shoots originate. Bromelioideae are generally terrestrial and Artigo recebido em 06/2007. Aceito para publicação em 01/2008. 1Part of the PhD thesis of the first author, Universidade Estadual Paulista, Campus de Rio Claro. 2Universidade Estadual Paulista, Instituto de Biociências, Departamento de Botânica, C.P. 199, 13506-900, Rio Claro, SP, Brasil. 3CNPq fellowship; financial support by FAPESP 4Author for correspondence: [email protected] 114 Proença, S. L. & Sajo, M. G. epiphytic and their leaf sheaths are usually As for the rhizomes, studies on the root wide and imbricate forming tanks where the anatomy of Bromeliaceae are rare. Krauss water and minerals accumulate to be (1949), Pita & Menezes (2002) and Segecin absorbed by scales and by the adventitious & Scatena (2004) related the rhizome roots growing there (Benzing & Burt 1970, anatomy of the species they studied to the Benzing 2000). In the terrestrial species growth habit of the plants and/or to the without tanks, as some Bromelia and environment where they grow. Ananas, the roots absorb water and The present study describes the nutrients directly from the soil (Benzing et morpho-anatomy of the rhizomes and roots al. 1976). Most Tillandsioideae are of epiphytic and terrestrial bromeliads from epiphytes and have a reduced or absent root the cerrado of São Paulo State, with the system and specialized scales covering the aim of pointing out particular features of leaves; in these plants called “atmospheric the Bromeliaceae as well as possible or extreme” the scales help in the hydric adaptive features found in the cerrado and nutritional balance; the roots, when vegetation. present, are mainly involved in fixation (Benzing & Burt 1970; Benzing 1973). Pitcairnioideae are usually terrestrial and MATERIAL AND METHODS rupicolous and grow in mesic and xeric The material was collected in areas of habitats; they present a well developed root cerrado in São Paulo State, Brazil and system for fixation and water and nutrient identified by the first author. Vouchers are absorption. The leaf scales in this subfamily deposited at the Rioclarense Herbarium are less developed and have little or no (HRCB) of the State University of São Paulo water absorption function (Pittendrigh 1948, (Table 1). Benzing & Burt 1970, Benzing et al. 1976). For the anatomical study at least two The rhizomes of Bromeliaceae are not representatives of each species were used. well known although Billings (1904) had Roots and rhizomes were fixed in FAA 50 described the rhizome of Tillandsia usneoides (Johansen 1940) for 48h and later preserved L. and Krauss (1948) had carefully studied the in ethanol 50%. Cross sections were made morphology and the anatomy of the rhizome by free hand in the apical and median of Ananas comosus (L.) Merr. Later, Segecin regions of both organs. The sections were and Scatena (2004) described the rhizomes of stained with Safranin and Astra blue some Tillandsia and interpreted them as a (Bukatsch 1972 modified by Kraus & Arduin possible adaptation to the epiphytic habit. 1997) and mounted in glycerin gelatin Kaiser The stems of monocot do not present (1880). For Tillandsia usneoides only the a cambium but many species posses a lateral stem was studied since this species does not apical meristem that determines the shape present roots when adult. of the plant. This meristem, called primary Sections of fresh material were used thickening meristem, is responsible for to test the presence/absence of phenolic thickening of the stem and for the formation compounds (Johansen 1940), starch of the adventitious roots and of the vascular (Johansen 1940), lignin (Sass 1951), lipids connection among the stem, roots and (Gerlach 1984) and to determine the crystals leaves. In some monocot genera, a chemical nature (Chamberlain 1932). secondary thickening meristem is also The photomicrographs were taken using present, which contributes to the formation an Olympus BX40 photomicroscope and a of the stem body (Rudall 1991). Leica MZ12 stereomicroscope. Rodriguésia 59 (1): 113-128. 2008 Rhizome and root anatomy of Bromeliaceae 115 Table 1 – List of studied species Subfamilies/Species Life forms Place of collect Voucher BROMELIOIDEAE Acanthostachys strobilacea Epiphyte Reserva Biológica S. L. Proença 201 (Baker) L.B.Sm. de Moji-Guaçu Aechmea bromeliifolia (Rudge) Baker Epiphyte Área de Proteção S. L. Proença 212 Ambiental de Corumbataí Ananas ananassoides (Baker) L.B.Sm. Terrestrial Estação Experimental S. L. Proença 192 de Itirapina Billbergia distachia (Vell.) Mez Epiphyte Reserva Biológica S. L. Proença 208 de Moji-Guaçu Billbergia porteana Brongn. Epiphyte Reserva Biológica S. L. Proença 198 de Moji-Guaçu Bromelia balansae Mez Terrestrial Estação Experimental S. L. Proença 197 de Itirapina PITCAIRNIOIDEAE Dyckia tuberosa (Vell.) Beer Terrestrial Universidade de S. L. Proença 213 São Paulo (USP), Pirassununga TILLANDSIOIDEAE Tillandsia loliacea Mart. Epiphyte Área de Proteção S. L. Proença 202 ex Schult. & Schult. f. Ambiental de Corumbataí Tillandsia pohliana Mez Epiphyte Reserva Biológica S. L. Proença 186 de Moji-GuaçuPratânia S. L. Proença 216 Tillandsia recurvata (L.) L. Epiphyte Estação Experimental de S. L. Proença 194 ItirapinaReserva Biológica S. L. Proença 209 de Moji-GuaçuPratânia S. L. Proença 215 Tillandsia tenuifolia L. Epiphyte Reserva Biológica S. L. Proença 210 de Moji-Guaçu Tillandsia tricholepis Baker Epiphyte Estação Experimental de S. L. Proença 195 ItirapinaÁrea de Proteção S. L. Proença 204 Ambiental de Corumbataí Tillandsia usneoides (L.) L. Epiphyte Estação Experimental de Itirapina S. L. Proença 193 Vriesea sp. Epiphyte Estação Experimental de Itirapina S. L. Proença 217 Rodriguésia 59 (1): 113-128. 2008 116 Proença, S. L. & Sajo, M. G. RESULTS Ananas ananassoides and Bromelia Rhizome balansae) and in the Pitcairnioideae D. Most of the studied species are epiphytic, tuberosa, the coating tissue is a stratified cork. except for the terrestrial Ananas The cortex, formed by rounded ananassoides, Dyckia tuberosa and Bromelia parenchymatous cells of variable sizes, balansae. The rhizomes are vertically or presents leaf traces (Figs. 1a, b, f; 2a-c) axillary horizontally positioned with their apical portion buds and besides idioblasts with calcium pointing upward. In the median and basal oxalate raphides (Figs. 1a, b; 2c). The regions of all rhizomes, axillary buds develop endodermis is recognized by the position of and originate new rhizomes that, in the
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