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Floral Anatomy and Micromorphology of Hyacinthoides Italica (L.) Rothm

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Floral Anatomy and Micromorphology of Hyacinthoides Italica (L.) Rothm ISSN 2226-3063 e-ISSN 2227-9555 Modern Phytomorphology 12: 81–93, 2018 https://doi.org/10.5281/zenodo.1313005 RESEARCH ARTICLE Floral anatomy and micromorphology of Hyacinthoides italica (L.) Rothm. A case of complete stachyospory in Asparagaceæ Julie Zalko *, Thierry Deroin ** Institut Systématique Evolution Biodiversité (ISYEB), Muséum national d'Histoire naturelle, CNRS, Sorbonne Université, EPHE, 57 rue Cuvier, CP 39, F-75231 Paris cedex 05, France; * [email protected], ** [email protected] Received: 16.03.2018 | Accepted: 11.07.2018 | Published: 16.07.2018 Abstract Floral vasculature and gynoecium architecture were studied in Hyacinthoides italica (L.) Rothm. Each locule shelters two ovules inserted basally and surprisingly supplied by axis bundles only, while lateral bundles of the carpels branch out into strands along the epidermises of septal nectaries. We brought then to the fore an unexpected trade-off between the supply of upper ovules in H. non-scripta and that of secretory tissues in the studied species. Moreover, a new pattern of septal nectary is described, with outer and inner cavities. Keywords: Hyacinthoides italica, Hyacinthaceæ, Asparagales, carpel, floral anatomy, trade-off, vertical zonality of gynoecium Introduction the British Isles and Netherlands to the north. Among them, H. italica (L.) Rothm. The genus Hyacinthoides Heist. ex Fabr., is well distinguished by its distribution previously nested in Hyacinthaceæ (Kubitzki area circumscribed to the maritime Alps of et al. 1998), is now placed in Asparagaceæ France and Italy. This species was previously (APG 2009; APG et al. 2016) and its phylogeny named Scilla italica L. and Endymion italicus was recently revised (Grundmann et al. (L.) Chouard. It is a perennial bulbous plant, 2010). It comprises 11 species, growing blossoming from March to May in shady around the western Mediterranean basin places (Coste & Flahault 1937), and often and Atlantic European regions as far as introduced for ornamental purposes. © The Author(s) 2018. Published by Andriy Novikov, State Natural History Museum NAS of Ukraine on behalf of Modern Phytomorphology. This is an open access article under the Creative Commons BY-NC-ND license (http://creativecommons.org/ licenses/by-nc-nd/4.0/) freely available on https://phytomorphology.org/ . 82 Zalko & Deroin The floral anatomy, especially gynoecial Succinct observations of H. italica vasculature, was studied for near species flowers visitors were made in the Botanical by Van Tieghem (1875). Gatin provided Garden of Paris on March the 28th and the additional data about pedicel and receptacle 1st of April 2017 in order to understand anatomy of Liliaceæ s.l., examined the workings of the different highlighted briefly H. italica (under Scilla italica) and structures. emphasized the occurrence of “glandes septales” i.e. septal nectaries between the carpels (Gatin 1920). Septal nectaries are the Results result of a postgenital and partial fusion of contiguous carpels (Van Tieghem 1875). Anthetical floral morphology Each individual of H. italica builds a single blue-purple racemose inflorescence, bearing Material and methods ca. 20 actinomorphous trimerous flowers, erect on long ascending pedicels, and A complete inflorescence of H. italica was axillate by two unequal bracts (Fig. 1 C). Each collected at May 16th 2013 in the Botanical flower comprises six tepals in two whorls, Garden at the Muséum national d’Histoire six stamens and three carpels (Fig. 1 A, B, E, naturelle in Paris (48° 50′ 36.521″ N, G–J). Tepals are fused together at their bases 2° 21′ 36.839″ E) and fixed by FAA (90 % and stamen filaments are adnate to them ethanol 70 %, 5 % formalin, 5 % acetic acid) along a short base (Fig. 1 A, B). Anthers are for 48 h, then preserved in a mixture of introrse. Ovary is externally very papillate – water, ethanol and glycerol (equal volumes). except for the short basal narrowed smooth Inflorescence structure, as well as those of region, which may be named gynophore six anthetical flowers (and gynoecium in two (Figs 1 D, F, 2 E–I and 4 H, I), – while it of them) were studied with a stereoscopic exhibits six depressions i.e. prints of the microscope (Fig. 1). Four flowers were stamens (Fig. 1 E). Ovary has three locules dehydrated through a t-butyl series and (Fig. 3 G), each with two anatropous ovules embedded in paraffin (melting point: 58– inserted at its base (Figs 1 D–F, 3 T, U and 60 °C, Gerlach 1984). Serial transverse and 4 H, I). Carpels are wholly fused in the lower longitudinal sections were cut at a thickness part of the ovary (where ovules occur), and of 15 µm by a rotary microtome Leitz 1512 partially above (Figs 1 F, 3 T, U and 4 H). (Germany), then stained by Astrablue In cross section, narrow clefts are observed [Chroma® 1B 163] 0.5 % aq. and Ziehl’s between the carpels, corresponding to outer fuchsine [RAL® 320490-1000] 10 % aq. All opening of septal nectaries (Figs 1 E, 3 A and slides were mounted in Eukitt [O. Kindler 4 C, E, F). GmbH® E0214]. Slides are kept in the plant histological collection of the Muséum Flower vascular anatomy national d’Histoire naturelle, Paris, under In the pedicel, the stele is ordered in 6 main the range references Zalko 1–4. bundles in two alternate triangles, whose Floral vasculature was reconstituted branching wholly provide above the perianth by drawings of the serial sections using a and androecium. Small intermediate camera lucida, and then by superimposing bundles, variable in number, are visible tracing papers on them. too (Fig. 2 A). Tepals (t) and stamens (e) are Modern Phytomorphology 12, 2018 Floral anatomy and micromorphology of Hyacinthoides italica 83 Figure 1. Morphological study of Hyacinthoides italica reproductive structures: A, B – anthetical flower (top and side views); C – top of the inflorescence; D – longitudinal-tangential section of gynoecium; E – transverse-median section of gynoecium at its basis; F – longitudinal-median section of gynoecium; G–J – floral buds just before anthesis, which were used for seriate paraffin sections. Modern Phytomorphology 12, 2018 84 Zalko & Deroin Figure 2. Transverse ascending sections of the flower (A–J) and gynoecium (K–Q) of Hyacinthoides italica, from pedicel to ovule insertion level: e – stamen; lc – lateral carpel bundle; mc – median carpel bundle; n – septal nectary (inner or outer); slc – synlateral carpel bundle; t – tepal. Dotted line delimits a central zone with small clear cells (E–J). Broken line is carpel epidermis at the center (P–Q). Arrows show structures quoted in the texte. Xylem in black, phloem left in white, all vasculature dashed when confused, secretory zones dotted (D–Q). Modern Phytomorphology 12, 2018 Floral anatomy and micromorphology of Hyacinthoides italica 85 Figure 3. Transverse ascending sections of the gynoecium of Hyacinthoides italica, from the ovule insertion level to stigma (A–R) and longitudinal sections of ovary (S–U): same abbreviations as on Fig. 2. Modern Phytomorphology 12, 2018 86 Zalko & Deroin supplied by one bundle each, their traces and 3 A–D), are more distinct at the level being fused in a common bundle. The of the ovule apices (Fig. 3 E–H), and wholly two organs break up from the gynoecium separate above (Fig. 3 I). So all carpels together (Fig. 2 F–H), then become distinct open in a single ovary locule (Figs 3 I–P (Fig. 2 I). Gynoecium is supplied by ca. 12 and 4 H–I), trilobate in cross section and unequal bundles (Fig. 2 G). Three bundles are gradually decreasing toward the stylar wider and well distinct at the gynophore level zone (Fig. 3 I–P). At the top of the ovary, the (Fig. 2 G,H, arrows), each of them branch last three secondary lateral carpel bundles off above in a median carpel bundle and wholly fused (Fig. 3 L, M, U), on each side of two mediolateral ones (Fig. 2 I, J, mc, mlc), the three clefts between the carpels and then all of them becoming peripheral inside the fade above (Fig. 3 M, N, arrows). As usual, ovary wall (Fig. 2 K), just below the locules. only median carpel bundles occur in the At the center of gynoecium (i.e. placenta) style (Fig. 3 N–Q), fading at the level where remain three synaxial bundles (Fig. 2 L, stylar canal is filled by secretory cells. arrows) and three synlateral carpel bundles Vascular architecture of the gynoecium (Fig. 2 M, slc). Each synaxial bundle divides of H. italica is sketched on Fig. 5 A. in two axial ones (a), which get closer to the ovules (Figs 2 O–Q, arrows, and 3 A), and at Flower micromorphology and gynoecium least supply them (Figs 3 B and 4 B, arrows). structure So the two ovules of a same carpel are fed Secretory tissues are recognized basally, by a single axial bundle, while each carpel between the tepals (Fig. 2 D–I), between synlateral bundle (slc) divides in two lateral stamens and gynoecium (Fig. 2 E–H) and ones (Fig. 2 M, N, lc), whose branches do not between tepals and stamens (Fig. 2 G, H). irrigate any ovule, but the inner part of the All these secretory regions stained deeply septal nectaries obviously outlined by the with fuchsine (Fig. 4 A). Outer clefts of the epidermises of the adjacent carpels (Figs 2 P ovary wall, related to septal nectaries (n), and 3 C). Lateral carpel bundles branch out are distinguished just below the level of the into five pairs of secondary strands which locules floor (Fig. 2 I) and their covering enter progressively the carpel walls, the first epidermis is secretory shortly above (Figs 2 J, pair diverges rather low (Fig. 2 M, arrows), 3 S and 6, NSE). A septal nectary is wholly four lateral bundles are seen above (Fig.
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