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Comparative Bark Anatomy of Root and Stem in Styrax Camporum (Styracaceae)

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Comparative Bark Anatomy of Root and Stem in Styrax Camporum (Styracaceae) IAWA Journal, Vol. 26 (4), 2005: 477–487 COMPARATIVE BARK ANATOMY OF ROOT AND STEM IN STYRAX CAMPORUM (STYRACACEAE) Silvia Rodrigues Machado1, Carmen Regina Marcati2, Berta Lange de Morretes3 & Veronica Angyalossy3 SUMMARY The bark of Styrax camporum Pohl (Styracaceae) differs anatomically in the root and stem. Roots have layered secondary phloem; short sieve tubes with simple, transverse or more or less inclined sieve plates; fibres in tangential bands; astrosclereids; wide rays, and a poorly developed periderm. Stems have non-layered secondary phloem; longer sieve tubes with compound, scalariform, inclined sieve plates; sclerified cells and brachysclereids; a developed periderm, and a non-persistent rhytidome. Prismatic crystals, starch grains, phenolic compounds and lipidic contents were observed in root and stem bark cells. The differences between the secondary phloem of root and stem are discussed. Key words: Bark anatomy, root, secondary phloem, stem, Styracaceae, Styrax camporum. INTRODUCTION Styracaceae occur in relatively warm parts of the world, such as the Mediterranean re- gion, eastern Asia, the Malay Archipelago and parts of North and South America, and one genus in tropical Africa. This family is best known by the indumentum of stellate or scale-like hairs and for its production of benzoin (gum benzoin) and storax (Metcalfe & Chalk 1950; Hutchinson 1973; Spongberg 1976). Styrax L. is by far the largest and most widespread of the 11 genera in the Styraca- ceae, with about 130 of the c. 160 species in the family. It has a widespread but disjunct distribution, occurring in the Americas, eastern Asia, and the Mediterranean region. Over half the species of the genus occur in South America, where they are distributed among a wide array of habitats, including the lowland and montane rain forest, sub- paramo, tepui scrub, restinga, rocky grasslands, and cerrados (Nakajima & Monteiro 1986; Fritsch 2001). Some Brazilian species of Styrax, for example S. ferrugineus and S. camporum, produce resin when the bark is wounded (Mors & Rizzini 1966). According to Metcalfe and Chalk (1950), the resin Storax and Gum Benzoin obtained 1) Departamento de Botânica, Instituto de Biociências, Universidade Estadual Paulista, Botucatu SP, CP 510, CEP 18618-000, Brazil [E-mail: [email protected]]. 2) Departamento de Recursos Naturais – Ciências Florestais, Universidade Estadual Paulista, Botucatu SP, CP 237, CEP 18603-970, Brazil. 3) Departamento de Botânica, Universidade de São Paulo, São Paulo SP, CP 11461, CEP 05421- 970, Brazil. Associate Editor: Alex Wiedenhoeft Downloaded from Brill.com10/06/2021 05:11:54PM via free access 478 IAWA Journal, Vol. 26 (4), 2005 Machado et al. — Bark anatomy of Styrax camporum 479 from Styrax officinale and S. benzoin, respectively, are secreted chiefly in pathological intercellular spaces of the phloem and wood of the stem. Styrax camporum Pohl, known as “estoraque” or “beijoeiro”, is a widely distributed small tree or shrub in the cerrado vegetation of south-eastern Brazil (Nakajima & Mon- teiro 1986). Cerrados, savanna-like ecosystems, are characterized by seasonal rainfall with dry cool winters (Franco 2002). In the dry season, fire is common in cerrados, as it is for most savanna ecosystems. The woody vegetation of cerrados is adapted to fire (Coutinho 1990), and the bark of S. camporum represents an effective protection for the cambium (Miranda et al. 2002). Although bark is of great importance both for resin production (Fahn 1979; Nair 1995) and as a protective barrier, very little information is available about bark anatomy in Styrax. General information about stem bark from several Styracaceae genera, including periderm origin and secondary phloem structure, were given by Metcalfe and Chalk (1950). In 1987, Cutler et al. described the bark anatomy of the roots of many species in their manual on root identification and included 4 species of Styrax. This paper describes the bark anatomy of root and stem of S. camporum and relates the results to the environmental conditions of the cerrado. MATERIALS AND METHODS Bark samples were obtained from adult plants of Styrax camporum from the cerrado near Botucatu, State of São Paulo, Brazil, (22° 55ʼ S, 48° 30ʼ W). The vouchers and sam- ples of root and stem wood of one specimen are deposited, respectively, in the Her- barium (SPF 42.588) and the Wood Collection (SPFw 302-root and 303-stem) of the Botany Department, University of São Paulo. Root samples were taken 1 m from the stem base, and have a diameter of 2 cm. Stem samples were taken below the first branch, at a distance of 50 cm from the base of the stem, and have a diameter of 5 cm. Bark samples were fixed in FAA 50, dehydrated in alcohol and subsamples were embedded in paraffin (Johansen 1940). Transverse, radial, and tangential sections (8–10 μm) were double-stained with safranin-fast-green and mounted in Permount synthetic medium. Some fixed, unembedded samples were cut on a sliding microtome and the sections (12–15 μm) stained with 1% aqueous solution of safranin and astra blue (Roeser 1972). Specimens from both root and stem were macerated in a mixture of equal volumes of acetic acid and hydrogen peroxide at 60 °C (Johansen 1940) for 12 to 24 hours. The material was stained with 1% aque- ous solution of safranin and astra blue and mounted in glycerine. Freehand sections of fresh material were treated with Sudan IV for identification of lipids and with ferric chloride for the identification of phenolic compounds (Jensen 1962). Bark descriptions follow Trockenbrodt (1990), Angyalossy-Alfonso and Richter (1991) and Richter et al. (1996). Quantitative data are based on 30 individual counts; the statistical requirements for minimum numbers of measurements were fulfilled: N = (t-value)2 . (sample variance) / (accuracy of 10% × sample mean)2, following Freese (1967) and Eckblad (1991). The numerical values given in the descriptions are expressed as minimum - mean - maximum. Downloaded from Brill.com10/06/2021 05:11:54PM via free access 478 IAWA Journal, Vol. 26 (4), 2005 Machado et al. — Bark anatomy of Styrax camporum 479 RESULTS Root General description — Bark reddish brown externally and dark reddish internally; surface smooth; thickness variable (0.2–0.4 cm) in relation to root diameter/age; bal- samic odor. Periderm: developed (Fig. 1 & 2), shallow and with straight course; phel- lem layered, composed of 3–10 layers of radially flattened thin-walled cells (Fig. 2); phelloderm composed of 1–3 rows of radially flattened thin-walled cells; oil contents were detected in phelloderm cells and phenolic contens in all periderm cells. Secondary phloem: regularly layered from cambium toward periphery in the transverse section; layers formed by multiseriate tangential bands of parenchyma and conducting cells (thin-walled sieve tube elements and companion cells) regularly alternating with tan- gential rows of thick-walled cells (sclerenchyma) (Fig. 1 & 3). These rows are crossed by phloem rays. Axial parenchyma: arranged in uniseriate tangential bands and with intercellular spaces in the innermost region (Fig. 4); cells become dilated towards out- ermost region of the phloem. Sieve tube elements: solitary or in groups of 4–5 (Fig. 4 & 14), interspersed with parenchyma cells and radially arranged; irregular in outline, about 26–33–46 μm in diameter and 105–297–462 μm in length; sieve plates hori- zontal to inclined, predominantly simple (Fig. 14 & 15); rarely compound sieve plates were observed with 3–6 sieve areas in a scalariform pattern; lateral sieve areas poorly developed. Rays: uniseriate (including locally biseriate rays) to 3–4 cells wide (Fig. 7); 32–48–73 μm in width, 10–16–26 cells and 305–636–976 μm in height; heterocellular with more than 4 rows of upright and /or square marginal cells; dilated towards the periphery due to anticlinal cell division and /or tangential cell expansion (Fig. 1 & 3); when in contact with fibres in the outer phloem, these cells accumulate phenolic com- pounds, develop a thick lignified secondary wall and show a pronounced intrusive growth (Fig. 6). Fibres: arranged in regularly spaced tangential bands (Fig. 1, 3 & 16); 1098–1311–1723 μm in length and 32 μm in diameter; fibre walls with short projections bearing crystalliferous cells (Fig. 17 & 18). Sclereids: astrosclereids (Fig. 19) occur near the phloem fibres and differentiate from parenchyma cells; these cells are filled with phenolic compounds before becoming sclerified and undergoing intrusive growth (Fig. 5). Inorganic contents: prismatic crystals isolated in chambered axial parenchyma cells and in square and/or upright subdivided ray cells (Fig. 6 & 7). Organic contents: starch grains in parenchyma cells; phenolic compounds in parenchyma cells (Fig. 5) and sclereids (Fig. 6); lipids in procumbent idioblastic ray cells (Fig. 7). Stem General description — Bark grey-brown externally and dark reddish internally; surface scaly; thickness variable (0.1–0.3 cm); balsamic odor. Periderm: well-developed and undulating (Fig. 8 & 9); phellem composed of 20–30 layers of radially flattened cells with U-shaped thickenings; phelloderm poorly developed, composed of 2–3 rows of thin-walled cells. Rhytidome non-persistent, encompassing 4–5 periderm layers and including parts of the collapsed phloem. Phenolic compounds in all cells of phellem and phelloderm. Lenticels present. Secondary phloem: non-layered; small groups of scler- Downloaded from Brill.com10/06/2021 05:11:54PM via free access 480 IAWA Journal, Vol. 26 (4), 2005 Machado et al. — Bark anatomy of Styrax camporum 481 enchyma cells interspersed with parenchyma cells (Fig. 8, 9 &10). Axial parenchyma: rectangular thin-walled cells, without intercellular spaces, in the innermost region of bark (Fig. 9); in the outer portion the parenchyma cells are tangentially enlarged and some cells become sclerified. Sieve tube elements: solitary or in groups of two, inter- spersed with parenchyma cells; about 25–27–30 μm in diameter and 340–390–480 μm in length; sieve plates inclined, compound, with 3–8 sieve areas in a scalariform pattern; lateral sieve areas prominent (Fig.
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