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Wood Anatomy of Family Salvadoraceae from the Indian Subcontinent with Special Reference to the Ultrastructure of the Vessel

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Wood Anatomy of Family Salvadoraceae from the Indian Subcontinent with Special Reference to the Ultrastructure of the Vessel Aliso: A Journal of Systematic and Evolutionary Botany Volume 29 | Issue 1 Article 8 2011 Wood Anatomy of Family Salvadoraceae from the Indian Subcontinent with Special Reference to the Ultrastructure of the Vessel Wall Vishakha Saxena Forest Research Institute, Dehradun, India Sangeeta Gupta Forest Research Institute, Dehradun, India Follow this and additional works at: http://scholarship.claremont.edu/aliso Part of the Botany Commons, and the Ecology and Evolutionary Biology Commons Recommended Citation Saxena, Vishakha and Gupta, Sangeeta (2011) "Wood Anatomy of Family Salvadoraceae from the Indian Subcontinent with Special Reference to the Ultrastructure of the Vessel Wall," Aliso: A Journal of Systematic and Evolutionary Botany: Vol. 29: Iss. 1, Article 8. Available at: http://scholarship.claremont.edu/aliso/vol29/iss1/8 Aliso, 29(1), pp. 59–63 ’ 2011, Rancho Santa Ana Botanic Garden WOOD ANATOMY OF FAMILY SALVADORACEAE FROM THE INDIAN SUBCONTINENT WITH SPECIAL REFERENCE TO THE ULTRASTRUCTURE OF THE VESSEL WALL VISHAKHA SAXENA AND SANGEETA GUPTA Wood Anatomy Discipline, Botany Division, Forest Research Institute, Dehradun, 248006, Uttarakhand, India ([email protected]) ABSTRACT The present study describes the wood microstructure of Azima tetracantha, Salvadora oleoides, and S. persica, the members of family Salvadoraceae represented in the Indian Subcontinent. An identification key based on wood anatomical features has been developed for the separation of the species. SEM studies revealed the presence of vesturing in intervessel pits of Salvadora. Key words: Azima, ecology, evolution, Salvadora, Salvadoraceae, vestured pits, wood anatomy. INTRODUCTION Saxena (2005) described perforated ray cells in order Celastrales. Family Salvadoraceae, the mustard-tree family, consists of three genera (Azima L., Dobera Juss., and Salvadora L.) with 12 species. It comprises trees, shrubs, and scramblers distri- MATERIAL AND METHODS buted in dry, hot regions of Africa, Madagascar, and Asia. The wood anatomical microstructure of Azima tetracantha, Two genera and three species are represented in the Indian Salvadora oleoides, and S. persica was studied as per the Subcontinent. Azima has two species of erect, thorny shrubs feature list developed by the International Association of distributed in Myanmar and India. Only one species, Azima Wood Anatomists (IAWA Committee 1989). The wood tetracantha Lam., occurs in India (Purkayastha 1985). samples featured in Table 1 are housed at the Xylarium of Salvadora has five species of shrubs and small trees distributed Forest Research Institute (DDw), Dehradun, India. in warmer parts of Africa and Asia, of which two—S. oleoides For microscopic examination and photomicrographs, 15– Decne. and S. persica L.—occur in India (Purkayastha 1985). 20 mm-thick transectional (Fig. 1–2), tangential (Fig. 3–4, 6), Salvadora occurs in a diverse array of habitats, ranging (in and radial (Fig. 5) sections were cut on a Reichert sliding order of increasing aridity) from the margins of mangroves, microtome, stained in Heidenhain’s hematoxylin and safranin, saline environments, swamps, and near riverbanks where the and mounted following a standard laboratory protocol, i.e., ground water level is high, to seasonally wet sites, grass passing through an ethanol (10–100%) and xylol series to savannah, thorn scrub, along irrigation lines in arid zones, and make permanent slides. For determination of fiber and vessel desert flood plains, indicating its tolerance of a wide range of element length, small radial chips were macerated following water quality, soil, and soil pH conditions (Zodape and Schultz’ method (30% nitric acid and a pinch of potassium Indusekhar 1997; Ahmad 2007). chlorate). Thirty measurements were taken for fiber length, The wood of Salvadora persica, the toothbrush tree (Almas vessel length, fiber diameter, and lumen diameter from 2002), has useful antiseptic properties, contributing to its use macerated material. The mean tangential diameter of the in the manufacture of toothpicks (Miswak sticks) for oral vessels was determined from 25 measurements from transverse hygiene. It is also used by the Egyptians to make coffins. The sections taken along the maximum width in radial direction. wood of S. oleoides is used for construction and in the The frequency of vessels was determined from the average of 10 manufacture of agricultural implements, Persian wheels counts per mm2 area. Ray frequency (rays per mm) was based (wheels operated by draught animals to lift water), and the on an average of 10 counts per linear mm in tangential sections. knees of boats. Almost all parts of Salvadora spp. have been Maximum ray height and ray width was also determined from found to be pharmaceutically important. The leaves, roots, tangential sections (Table 2). Data on specific gravity and gross bark, fruits, and seeds are used for the treatment of cough, features follow Purkayastha (1985). Ecological adaptability was fever, asthma, and as a purgative. Roots are also used to treat measured in terms of vulnerability (mean vessel diameter/mean chest infections and the latex to treat sores (Mahar and Malik vessel frequency) and mesomorphy (vulnerability 3 mean vessel 2001; Savithramma et al. 2007). length) of each sample. Intervessel pitting was studied by Prior to this work, no detailed account of microscopic wood scanning electron microscopy. structure was available for family Salvadoraceae from India. Gamble (1922) described gross features of these species in RESULTS brief. Metcalfe and Chalk (1950) summarized microscopic features at the generic level. Den Outer and Van Veenendaal Generic Description and Identification Key (1981) described wood and bark anatomy of Azima tetra- Azima cantha, and Carlquist (2002) provided some detail of wood and bark anatomy of the family. Saxena and Gupta (2004) Azima tetracantha.—Growth rings indistinct. Wood diffuse reported perforated ray cells in Salvadoraceae, and Gupta and porous, vessels exclusively solitary, rarely clustered, round to 60 Saxena and Gupta ALISO Table 1. Species of Salvadoraceae studied, with information on accession number, collection locality, specific gravity, vulnerability, and mesomorphy. Species Accession number Locality Specific gravity Vulnerability Mesomorphy Azima tetracantha DDw 5639 India: Tamil Nadu State: Coimbatore 0.63 1.6 428 Salvadora oleoides DDw 1382 Pakistan: Sindh Province 0.85 6.1 831 Salvadora persica DDw 1381 Pakistan: Sindh Province 0.49 5.5 935 DDw 5129 Pakistan: Sindh Province 0.58 8.0 1456 DDw 5695 India: Tamil Nadu State: Coimbatore 0.67 2.9 647 oval. Perforation plates simple. Mean tangential diameter of DISCUSSION vessel 72 (19–130) mm, mean vessel element length 262 (161– Carlquist (2002) reported anatomical characteristics of 375) mm. Intervessel pits non-vestured, alternate, 3 mmin Azima tetracantha and Salvadora persica from South Africa, diameter, vessel–ray pits similar to intervessel pits in shape and and in Table 3 we compare our data to his. Wide variation was size. Vessel frequency 33–55/mm2. Fibers non-septate, mean observed in the quantitative anatomical features. These fiber length 817 (642–1017) mm, mean fiber wall thickness differences likely can be attributed to site conditions, age of 10 mm, thick-walled with slit-like apertures, mean fiber the plant, or growth form. diameter 17 (14–22) mm. Parenchyma storied, in wavy While studying the wood microstructure, we observed wall bands up to 3–12 cells wide, in strands of 3–4 cells, strands Salvadora sometimes fusiform. Rays 1–21-seriate (Fig. 4), of two distinct ingrowths in the minute intervessel pits of . Scanning sizes, heterogeneous with mostly procumbent, square, and electron microscopy showed these ingrowths to be vestures upright cells mixed throughout the rays. Ray frequency 1–4/ in the intervessel pits. The occurrence of vestured pits in Salvadora is consistent with the prevalence of pit vesturing mm, mean uniseriate ray height 214 (107–331) mm, 3–12- in plants from tropical environments (Jansen et al. 2004). celled, mean uniseriate ray width 11 (8–14) mm, mean The present study is the first to report vestures in genus multiseriate ray height 890 (268–3467) mm, 9–14-celled, mean Salvadora. multiseriate ray width 142 (32–407) mm. Perforated ray cells present (Fig. 2). Prismatic non-chambered crystals present in Vulnerability and mesomorphy were proposed by Carlquist ray cells. (1977) as indices of ecological variation in wood anatomy. Low values of vulnerability (,1.0) and mesomorphy (,75) Salvadora indicate adaptation to xeric conditions, i.e., the capacity of species to resist water or freezing stress, whereas high values of Salvadora oleoides and S. persica.—Growth rings indistinct. vulnerability (1.0–2.5) and mesomorphy (.200) show adapta- Wood diffuse porous, vessels solitary and in radial multiples of tion to mesic conditions. Consistent with their tropical origin, 2–5 cells, few in clusters, round to oval. Perforation plates our samples show vulnerability values of .1 and mesomorphy simple. Mean tangential diameter of vessels 61 (22–122) mm, values of .200. mean vessel element length 178 (75–268) mm. Intervessel pits Several wood characters are seen as indicators of phyloge- vestured, alternate, 3–5 mm in diameter (Fig. 5–6), vessel–ray netic advancement, i.e., short vessel elements, simple perfora- pits similar to intervessel pits in shape and size. Vessel tion, minutely bordered pits, storied features, and the ratio of frequency 5–21/mm2. Fibers non-septate
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