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Structural and Histochemical Characterization of the Osmophores in Corollas of Asteraceae (Tribes Onoserideae and Famatinantheae)

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Structural and Histochemical Characterization of the Osmophores in Corollas of Asteraceae (Tribes Onoserideae and Famatinantheae) adansonia 2020 42 12 DIRECTEUR DE LA PUBLICATION / PUBLICATION DIRECTOR: Bruno David Président du Muséum national d’Histoire naturelle RÉDACTEUR EN CHEF / EDITOR-IN-CHIEF : Thierry Deroin RÉDACTEURS / EDITORS : Porter P. Lowry II ; Zachary S. Rogers ASSISTANT DE RÉDACTION / ASSISTANT EDITOR : Emmanuel Côtez ([email protected]) MISE EN PAGE / PAGE LAYOUT : Emmanuel Côtez COMITÉ SCIENTIFIQUE / SCIENTIFIC BOARD : P. Baas (Nationaal Herbarium Nederland, Wageningen) F. Blasco (CNRS, Toulouse) M. W. Callmander (Conservatoire et Jardin botaniques de la Ville de Genève) J. A. Doyle (University of California, Davis) P. K. Endress (Institute of Systematic Botany, Zürich) P. Feldmann (Cirad, Montpellier) L. Gautier (Conservatoire et Jardins botaniques de la Ville de Genève) F. Ghahremaninejad (Kharazmi University, Téhéran) K. Iwatsuki (Museum of Nature and Human Activities, Hyogo) K. Kubitzki (Institut für Allgemeine Botanik, Hamburg) J.-Y. Lesouef (Conservatoire botanique de Brest) P. Morat (Muséum national d’Histoire naturelle, Paris) J. Munzinger (Institut de Recherche pour le Développement, Montpellier) S. E. Rakotoarisoa (Millenium Seed Bank, Royal Botanic Gardens Kew, Madagascar Conservation Centre, Antananarivo) É. A. Rakotobe (Centre d’Applications des Recherches pharmaceutiques, Antananarivo) P. H. Raven (Missouri Botanical Garden, St. Louis) G. Tohmé (Conseil national de la Recherche scientifique Liban, Beyrouth) J. G. West (Australian National Herbarium, Canberra) J. R. Wood (Oxford) COUVERTURE / COVER : Réalisée à partir des Figures de l’article/Made from the Figures of the article. Adansonia est indexé dans / Adansonia is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Agriculture, Biology, and Environmental Sciences® – Scopus® Adansonia est distribué en version électronique par / Adansonia is distributed electronically by: – BioOne® (http://www.bioone.org) Adansonia est une revue en flux continu publiée par les Publications scientifiquesdu Muséum, Paris Adansonia is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Geodiversitas, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie, Comptes Rendus Palevol Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2020 ISSN (imprimé / print) : 1280-8571/ ISSN (électronique / electronic) : 1639-4798 Structural and histochemical characterization of the osmophores in corollas of Asteraceae (tribes Onoserideae and Famatinantheae) Liliana KATINAS División PlantasVasculares, Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata (Argentina) [email protected] (corresponding author) Marcelo P. HERNÁNDEZ División PlantasVasculares, Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata (Argentina) and Facultad de Ciencias Agrarias y Forestales, 60 y 119, Universidad Nacional de La Plata, 1900 La Plata (Argentina) Gisela SANCHO División PlantasVasculares, Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata (Argentina) Submitted on 14 August 2019 | accepted on 24 June 2020 | published on 14 september 2020 Katinas L., Hernández M. P. & Sancho G. 2020. — Structural and histochemical characterization of the osmophores in corollas of Asteraceae (tribes Onoserideae and Famatinantheae). Adansonia, sér. 3, 42 (12): 227-239. https://doi. org/10.5252/adansonia2020v42a12. http://adansonia.com/42/12 ABSTRACT Osmophores are engaged in scent production and differ from other secretory structures by their product, site, duration, and anatomical structure. Whereas osmophores have been well-documented in flowers of several families, they are barely mentioned in the Asteraceae. The aims of this study are to: 1) determine the occurrence of osmophores in corollas of 39 species of the tribe Onoserideae and the only species of Famatinantheae with histochemical methods; and 2) analyze the morphology and structure of osmophores in these groups. Histochemical and histological techniques revealed osmophores in the marginal and in the central corollas of Famatinanthus Ariza & S.E. Freire and Plazia Ruiz & Pav. at the apex and margin of the corolla lobes, and at the sinuses. Osmophores KEY WORDS were confirmed following positive staining reactions for TIOFH and TIOFH3 (Neutral Red, Oil Anatomy, Asteraceae, Red O, and Iodine-Potassium-Iodide); the samples reacted negatively for Benedict’s test for detect- Compositae, ing reducing sugars. The osmophores found in this study are constituted uniquely by a papillose or histochemistry, non-papillose epidermis, or they extend into the mesophyll forming scent glands. In both cases, they osmophore, scent production, are associated to stomata and starch grains. The corollas of the other species of Onoserideae did not secretory structures. react positively for osmophores. ADANSONIA, sér. 3 • 2020 • 42 (12) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.adansonia.com 227 Katinas L. et al. RÉSUMÉ Caractérisation structurale et histochimique des osmophores dans les corolles d’Asteraceae (tribus des Ono- serideae et des Famatinantheae). Les osmophores participent à la production d’odeurs et se différencient des autres structures de sécrétion par leur produit, leur localisation, leur durée et leur structure anatomique. Alors que les osmophores ont été bien documentés dans les fleurs de plusieurs familles, ils sont à peine mentionnés dans les Asteraceae. Les objectifs de cette étude sont les suivants : 1) déterminer la présence d’osmophores dans les corolles de 39 espèces de la tribu des Onoserideae et de la seule espèce de Famatinantheae en utilisant des méthodes histochimiques ; et 2) analyser la morphologie et la structure des osmophores dans ces groupes. Les techniques histochimiques et histologiques ont révélé la présence d’osmophores dans les corolles marginales et centrales de Famatinanthus Ariza & S.E. Freire and Plazia Ruiz & Pav. au sommet et au bord des lobes de la corolle et aux sinus. Les osmophores ont été confirmés à la suite MOTS CLÉS de réactions de coloration positives pour TIOFH et TIOFH3 (Rouge Neutre, Oil Red O et Iodure Anatomie, Asteraceae, de Potassium Iodé) ; les échantillons ont réagi négativement au test de Benedict pour la détection Compositae, des sucres réducteurs. Les osmophores trouvés dans cette étude sont constitués uniquement d’un histochimie, épiderme papilleux ou non, ou s’étendent dans le mésophylle formant les glandes odorantes. Dans osmophore, production de parfum, les deux cas, ils sont associés à des stomates et à des grains d’amidon. Les corolles des autres espèces structures de sécrétion. d’Onoserideae n’ont pas réagi positivement pour les osmophores. INTRODUCTION why osmophores are barely mentioned in the family. Field observations and herbarium label information, however, show Osmophores parts are engaged in scent production and differ that many members of this cosmopolitan family have, or are from other secretory structures (e.g., hydathodes, nectaries, resin described with, fragrant flowers. Detailed studies of osmophores ducts) by their product, site, duration, and anatomical structure, in Asteraceae are lacking; they are only briefly mentioned, for and they are commonly found in certain floral and inflorescence example, as a complement of floral pollination studies. In a parts (Stern et al. 1987; Vogel 1990; Effmertet al. 2005; Wie- study of reproductive strategies of Mikania glomerata Spreng. mer et al. 2009). The osmophoric function may be fulfilled only and M. hirsutissima DC. (Eupatorieae), Eiterer (1965) rejects by the epidermis (glandular epithelium), sometimes papillose the presence of osmophores in these taxa arguing that the or with trichomes, or the glandular structure may also extend odor is produced by the pollen grains, the anthers, and the into the mesophyll tissue forming scent glands (Vogel 1990). stigmatic lines of the styles. Combas et al. (1999), in a floral The fragrant substances secreted by osmophores are mainly biology study, localize osmophores in the veins, margins and lipophilic, volatile, low terpenes, which are the primary con- base of the neutral corollas of Tithonia diversifolia (Hemsl.) stituents of the essential oils. They occur in the form of minute A. Gray (Heliantheae). Santos de Castro et al. (2015) indicate droplets in the cytoplasm of the epidermal and neighboring that the odor in florets of Moquiniastrum oligocephalum (Gard- parenchyma cells. At an appropriate temperature, the droplets ner) G. Sancho (Gochnatieae) is due to osmophores located diffuse in gaseous form from the cytoplasm through the cell in the corolla lobes, styles and anther apices. Melo Santos wall and cuticle to the outside. With the diffusion of droplets, et al. (2016) mention osmophores in Tridax procumbens L. new ones are constantly produced. Below the epidermis the cells (Heliantheae) with an unclear location, although indicating are filled with starch grains which, used as a source of energy, petals and ‘sepals’ as possible scent-producing sites. reveal the secretory activity of the osmophores (Fahn 1979). During a routine light microscope examination of florets Osmophores have been
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