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Vanessa Koza Kowalski1,2, Pamella Paula Diniz Alves Pereira1, Fernanda Maria Cordeiro De Oliveira1, Maria Eugênia Costa1 & Rosangela Capuano Tardivo1

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Vanessa Koza Kowalski1,2, Pamella Paula Diniz Alves Pereira1, Fernanda Maria Cordeiro De Oliveira1, Maria Eugênia Costa1 & Rosangela Capuano Tardivo1 Rodriguésia 67(2): 427-435. 2016 http://rodriguesia.jbrj.gov.br DOI: 10.1590/2175-7860201667213 Are the wing’s cells alive? Study case in Vriesea trichomes As células da ala são vivas? Estudo de caso em tricomas de Vriesea Vanessa Koza Kowalski1,2, Pamella Paula Diniz Alves Pereira1, Fernanda Maria Cordeiro de Oliveira1, Maria Eugênia Costa1 & Rosangela Capuano Tardivo1 Abstract The presence of peltate foliar trichomes is one of the main anatomical characteristic of Bromeliaceae. These complex structures are adapted to compensate water and nutrient absorption in species that have reduced or substrate and light-reflection independent roots. They have enabled species’ survival in diverse and extreme environments contributing to the wide distribution of this family. In the present work, we analyzed the peltate trichomes’ characteristics in three taxa of Vriesea (Tillandsioideae): Vriesea platynema var. platynema, V. platynema var. variegata and V. tijucana. Leaves in different developmental stages were analyzed with histochemical tests and Transmission Electron Microscopy. Main results include the presence of cytoplasmic content in the wing peripheral cells, as well as in mature leaves. This is the first register of the presence of such feature in this family, which brings the possibility of discussing how water can be absorbed by these cells. Key words: Anatomy, Tillandsioideae, absorptive trichomes. Resumo Uma das principais características anatômicas de Bromeliaceae é a presença de tricomas foliares peltados. Estas complexas estruturas adaptadas à compensação da absorção de água e alimento em espécies com raízes reduzidas ou independentes do substrato e reflexão da luz, possibilitaram a vida em ambientes diversos e extremos, contribuindo para a ampla distribuição desta família. Neste estudo são analisadas as características dos tricomas peltados de três espécies do gênero Vriesea (Tillandsioideae): Vriesea platynema var. platynema, V. platynema var. variegata e V. tijucana. Folhas em diferentes fases de desenvolvimento foram analisadas com testes histoquímicos e de Microscopia Eletrônica de Transmissão. Destacam-se a presença de citoplasma evidente em células periféricas da ala, inclusive em folhas maduras, característica esta, inédita evidenciada pela primeira vez para a família, possibilitando a discussão da absorção de água por estas células. Palavras-chave: Anatomia, Tillandsioideae, tricomas absorventes. Introduction leaves have enabled survival in diverse and extreme Bromeliaceae Juss. is one of the most striking environments, contributing to the wide distribution families in the tropical forests of the American of the family (Benzing 2000). continent (Benzing 2000; Martinelli et al. 2008). Hypotheses on the evolution of epiphytism It comprises 58 genera and 3,352 currently known in Bromeliaceae, which is mainly observed in species (Luther 2012), which are distributed in Tillandsioideae have been put forth by a number of three major regions: the Atlantic Forest along the authors. According to Schimper (1888), epiphytes eastern coast of Brazil, the Guyana Highlands, evolved from terrestrial species found in rainforests and from the northern Andes to Mexico and the growing apart from their usual source of water, the Antilles. Evolutionary aspects, such as leaves soil. Tietze (1906) states that the likely ancestor arranged in a rosette enabling rainwater capture of Bromeliaceae would be a xerophytic plant, and storage, as well as the absorptive trichomes in similar to the species of Pitcairnioideae. This 1 Universidade Estadual de Ponta Grossa, Campus Uvaranas, Av. General Carlos Cavalcanti 4748, 84030-900, Ponta Grossa, PR, Brazil. 2 Author for correspondence: [email protected] 428 Kowalski, V.K. et al. subfamily has foliar trichomes that potentially have also described the basal stalk and the dome- perform the function of absorbing water given cells with densely cytoplasmic contents. Recently, their contact with the mesophyll, despite a low studies elaborated by Proença & Sajo (2004, 2007); degree of organization. In contrast, trichomes Mantovani & Iglesias (2005); Scatena & Segecin in Bromelioideae and Tillandsioideae have a (2005); Souza, Estelita & Wanderley (2005); greater degree of organization and constitute an Pereira et al. (2011); Silva & Scatena (2011); efficient water and nutrient absorption system. The and Silva, Oliveira & Scatena (2011) described hypothesis proposed by Pittendrigh (1948) is in the trichome structure in different genera of partial agreement with the one proposed by Tietze Bromeliaceae, but made no reference to either the (1906), in which the likely ancestor is a terrestrial presence or absence of content within the cells. plant from dry environments. However, the Considering the works mentioned above, ascendance to Pitcairnioideae suggested by Tietze the present study aimed to describe the structure (1906) is doubtful. Pittendrigh (1948) suggests that of peltate trichomes in three taxa of Vriesea Lindl. formation of the leaf rosette is involved in water (Tillandsioideae): Vriesea platynema Gaudich. and humus capturing through the leaves’ bases and var. platynema, V. platynema var. variegata states that the absorbent foliar trichome system (Guillon) Reitz, and V. tijucana E. Pereira, offering has enabled the existence of epiphytic species in a discussion on the cytoplasmic content in the Bromeliaceae. wings’ peripheral cells of the peltate trichomes in Anatomically, Bromeliaceae has typical Bromeliaceae. characteristics such as: epidermal cells with sinuous walls containing silica bodies (Smith & Downs Material and Methods 1974; Tomlinson 1969), and peltate trichomes on the leaves which are adapted to water and nutrient The taxa analyzed here are all epiphytic absorption in species with short roots and roots that ones. They bear their leaves arranged in a rosette do not depend on substrate or light reflection. The that forms a basal tank for capturing water and latter is considered to be an apomorphic feature for nutrients. Sampling was carried out at different the family (Gilmartin & Brown 1987). sites in Paraná State (Brazil), comprising regions of The pedicel connects the trichomes to the mixed rainforest, dense rainforest and grasslands. remaining epidermis and the wing in order to Part of the sampled material was deposited at the capture and transport water and nutrients. The Herbarium of Universidade Estadual de Ponta wing is positioned in parallel to the leaf surface Grossa (Brazil) under the following numbers: and is composed of a central disc and a peripheral Vriesea platynema Gaudich. var. platynema (V. wing, which vary in number and position according Kowalski 22, 58, 59), Vriesea platynema var. to each subfamily in response to the evolutional variegata (Guillon) Reitz (V. Kowalski 09, 28, pattern (Strehl 1983). 34), and Vriesea tijucana E. Pereira (V. Kowalski Anatomical studies in Bromeliaceae have 17, 48, 55). The remaining material was fixed in been carried out with different goals, such formaldehyde-ethanol-acetic acid 50% (FAA50) for as the characterization of its subfamilies, the 24 hours (Johansen 1940) and stored in 70% ethyl understanding of the plant-environment relationship alcohol. In addition, living plants were cultivated and taxonomic classifications. for histochemical tests. Specific studies addressing absorbent peltate Three plants of each species were selected trichomes have been conducted since the 19th from different sites as a means to ensure the century traditionally analyzing three subfamilies: analysis of individuals with different genotypes. Bromelioideae, Pitcairnioideae and Tillandsioideae. According to their position within the rosette, Mez (1896) accounts the structure of trichomes and young, mature and old leaves were selected and Krauss (1949) detailed their ontogeny, but neither sectioned at the half portion of the leaf blade and of them pointed out details of cell’s content. also at the half portion of the leaf sheath. Tomlinson (1969), Benzing (1976), Braga (1977) Paradermal, transverse and longitudinal and Strehl (1983) have detailed the structure, the sections were produced by free hand sectioning number and the shape of cells. These authors have using razor blades. The sections were stained with described the structure of wing cells, also known safranin, basic fuchsine, Astra blue and Toluidine as shield cells, to be dead, losing cell content when Blue (Kraus & Arduin 1997). Glycerinated gelatin trichomes reach their full developed. These authors was used as the mounting medium (Kaiser 1880). Rodriguésia 67(2): 427-435. 2016 Cytoplasmic content in Wing’s cells of Trichomes 429 The sections were submitted to histochemical Ultracut S microtome and stained with Toluidine analysis with phloroglucin solution in an acid Blue (O’Brien et al. 1964). Ultrathin sections were medium for the detection of lignin (Berlyn & stained with uranyl acetate (Gibbons & Grimstone Miksche 1976). Parts of fresh samples were 1960) followed by lead citrate (Reynolds 1963). submitted to the tetrazolium test in order to The material was analyzed using Zeiss EM900 determine the vitality of the peltate trichomes’ wing microscope at the Microscopy Center of the cells. The tetrazolium test indicates the presence University of São Paulo. of living tissue by binding to respiration enzymes inside these cells. When in contact with tetrazolium, Results these enzymes produce trifenilformazan, which Although we analyzed three different stages gets colored in red (Brasil
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