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Taxonomic Relevance of Calcium Oxalate Cuticular Deposits in Dracaena Vand. Ex L

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Taxonomic Relevance of Calcium Oxalate Cuticular Deposits in Dracaena Vand. Ex L HORTSCIENCE 36(6):1033–1036. 2001. lar crystals, but members of Sansevieria ex- hibit minute (<1 mm) crystals in the cuticle (Pennisi, 1999). The objectives of this study Taxonomic Relevance of Calcium were to investigate the occurrence of crystalliferous cuticle in selected Dracaena Oxalate Cuticular Deposits in species in order to determine its taxonomic relevance in providing additional characters Dracaena Vand. ex L. for segregation of cultivated Dracaena species. Svoboda V. Pennisi1 Materials and Methods Department of Horticulture, University of Georgia, Coastal Plains Experiment Plant materials. The Dracaena species Station, Tifton, GA 31793 used in this study were obtained from two 2 general sources. The cultivated group included Dennis B. McConnell those species commonly grown as foliage Department of Environmental Horticulture, University of Florida, Gainesville, plants in the United States. The second group FL 32611-0670 was the dragon tree group from Spain and Gran Canaria, although D. draco (L.) L. is Additional index words. crystalliferous cuticle, cuticular periplasmic crystals, Dracaenaceae, commonly grown in the United States. The Agavaceae, Cordyline, Sansevieria following species were compared: Dracaena arborea (Willd.) Link; D. cincta; D. deremensis Abstract. Detection of cuticular crystals in the 14 species of Dracaena examined indicated Engl.; D. draco; D. fragrans; Dracaena that they are probably ubiquitous throughout the genus and may permit rapid separation ×massefiana hort. (a hybrid between D. of dracaenas from plants with similar leaves such as the cordylines (Cordyline sp.). fragrans ‘Massangeana’ and D. surculosa); Dracaena species of the dragon tree group deposit the greatest quantity of uniformly small D. reflexa (Descne.) Lam.; D. sanderiana; D. cuticular crystals. However, the distinction between individual species within this group- surculosa Lindl.; D. thalioides hort. Makoy ing, based solely on crystal numbers and size, is not sufficient for taxonomic separation. ex E. Morr.; Cordyline australis (Forst.) End.; All other species of Dracaena studied did display species-specific quantities and sizes of C. terminalis (L.) Kunth; Sansevieria cuticular crystals. This, in combination with characteristics of the leaf epidermis, could cylindrica Bojer.; and S. trifasciata hort. ex serve as part of a taxonomic key to the genus. Prain. All Cordyline, Dracaena, and Sansevi- eria species were grown in the conservatory of The monocotyledonous genus Dracaena Most of the cultivars commercially grown the Environmental Horticulture Dept. at Vand. ex L. consists of ≈60 species have variegated leaves. Within the genus, Gainesville except D. arborea, which was (Hutchinson, 1986), and all members except leaves are variable in length (≈40 cm long) and grown at a local nursery. In addition, her- one (D. americana J.D. Sm.) are indigenous to are narrow (<1.5 cm) to broad (>5 cm). barium specimen of four species from the tropical regions of Africa and Asia. Cultivated Hutchinson (1959) assigned Dracaena to Jardin Botanico Canaria “Viera y Claijo” (JBC) species of Dracaena are popular houseplants Agavaceae Endl., but Takhtajan (1980) placed in Spain were examined: D. cinnabari Balf.; and widely used in interior landscapes. Also, it in a separate family, Dracaenaceae Salisb., D. ellenbeckiana Engl.; D. ombet Kots. & Dracaena is the most economically important related to Asparagaceae Juss. (or Liliaceae Peyr.; and D. tamaranae Marrero et al. The genus of foliage plants grown in Florida Juss. subfam. Asparagoideae Kostel). Recent first three species had been grown in the JBC, (McConnell et al., 1989). They are usually analysis of internal transcribed spacer (ITS) while the last species originated from Gran sold as small juvenile plants (20 cm to 2 m in rDNA sequence of 40 taxa in Agavaceae jus- Canaria. Authorities for these herbarium speci- height), but in their native habitats, some spe- tified this placement in Dracaenaceae (Bogler mens are as listed in Marrero et al. (1998). All cies reach considerable heights (6 to 20 m). and Simpson, 1996). Sansevieria Thunb. forms specimens from Spain, along with D. draco, Because the juvenile stage is commonly used, a monophyletic group with Dracaena and the are considered to be in the dragon tree group, identification is based primarily on leaf mor- genera are very closely related. The vegetative an arborescent taxa of Dracaena found in East phology. Leaves of Dracaena species may be and floral anatomy of some species of and West Africa (Marrero et al., 1998). pliable or stiff and their shape varies from Cordyline Comm. ex R. Br. closely resemble Methods. Epidermal peels were obtained linear to sword-shaped to lanceolate, with those of some members of Dracaena (Bogler from leaves from three plants of each species clasping flaring leaf bases or distinct petioles. and Simpson, 1996). This similarity is often a using the following procedure. Fresh mature source of confusion in field identification, and leaf material (1 g) was cut into 10 × 10-mm Received for publication 18 May 2000. Accepted retail nurseries often sell species of Cordyline pieces and placed for 48 h in a maceration for publication 13 Dec. 2000. Florida Agricultural as red dracaenas. However, according to Bogler solution (10 mL) containing cellulase (1.0% Experimental Station journal series no. R-07510. and Simpson (1996), Cordyline has a clearly w/v), hemicellulase (1.0% w/v), and pectinase Based on part of a dissertation accepted in partial divergent ITS sequence, setting it apart from (0.1% w/v) (Protoplast Isolation Enzyme So- fulfillment of the requirements for a PhD degree at members of Dracaenaceae. lution I; Sigma-Aldrich Co., St. Louis). The the Univ. of Florida. Mention of a trademark, propri- Extracellular epidermal crystals in dracae- epidermal peels from herbarium specimens etary product, or vendor does not constitute a guar- antee or warranty of the product by the U.S. Dept. of nas were documented by Kohl (1889), who (two leaves from each species) were obtained Agriculture and does not imply its approval to the described relatively large rhombohedral crys- by soaking the leaves for 3 h in boiling water exclusion of other products or vendors that may also tals associated with epidermal cells of Dra- prior to enzymatic digestion. After macera- be suitable. The authors are grateful to Aguedo caena fragrans (L.) Ker-Gawl. Fink (1991) tion, adaxial and abaxial epidermal peels were Marrero and Rafael Almeida for herbarium samples reported that D. cincta Bak. (syn. D. marginata obtained by gently pulling the epidermis away and to Bijan Dehgan and Jake Henny for review of Lam.) had small crystals embedded within the from the underlying mesophyll. The peels the manuscript and instructive criticism. The cost of cuticular layer above the striated epidermal were rinsed in ethanol, placed on glass slides publishing this paper was defrayed in part by the cell wall. Pennisi et al. (2001) reported small and examined with a Nikon Optiphot-Pol re- payment of page charges. Under postal regulations, (<1 to 6 µm) crystals beneath the cuticle of search microscope (Nikon Nippon Kogaku this paper therefore must be hereby marked adver- tisement solely to indicate this fact. Dracaena sanderiana hort. Sander ex K.K., Tokyo) equipped with polarizing optics. 1Assistant Professor. M.T.Mast. These consisted of calcium oxalate Detailed cellular measurements were made 2Professor. To whom requests for reprints should be monohydrate (COM) and were formed in close with an ocular micrometer and crystal counts addressed. E-mail address: dmcconnell@mail. proximity to the epidermal cell cytoplasm. were taken from micrographs. Photographs ifas.ufl.edu Members of Cordyline do not possess cuticu- were taken with an automatic Nikon UFX-II camera attachment (Nikon Nippon Kogaku smallest length to width ratio, 1.2:1. Exclud- that lead to deposition of calcium oxalate in K.K.). ing D. fragrans, the other species studied cuticular leaf areas. Conversely, these same possessed epidermal cells with length to width features are not shared among Dracaena-like Results and Discussion ratios of about 3:1 to 15:1. The orientation of Cordyline species. In terms of relationships crystal deposits was random in species that among genera, our findings agree with those All Dracaena species examined possessed displayed numerous minute cuticular crystals, of Bogler and Simpson (1996). cuticular crystals (Table 1). Cuticular deposits but less so in species that possessed larger, but Calcium oxalate (CO) crystals occur in in the cultivated group varied among Dra- fewer crystals. Dracaena fragrans was an more than 200 families of Magnoliophyta caena species with respect to quantity and size exception to these general trends, as it had Cronquist (syn. Angiospermae A. Braun & (Fig. 1). Within the dragon tree group charac- large but randomly distributed crystals. Crys- Doell) (Zindler-Frank, 1976) and Pinophyta teristics were almost identical with respect to tal orientation was related to epidermal cell Cronquist (syn. Gymnospermae Lindley) epidermal cell dimensions and crystal deposi- shape; the greater the length to width ratio and (Fink, 1991). Angiospermae are characterized tion (Fig. 1 A, D, and K–N). the larger the deposits, the more regular the by intracellular CO deposition, while a Among the cultivated species, D. thalioides orientation of the crystals. This was readily “crystalliferous cuticle” [extracellular crys- had the largest cuticular crystals (Fig.
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