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Bark Anatomy and Cell Size Variation in Quercus Faginea

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Bark Anatomy and Cell Size Variation in Quercus Faginea Turkish Journal of Botany Turk J Bot (2013) 37: 561-570 http://journals.tubitak.gov.tr/botany/ © TÜBİTAK Research Article doi:10.3906/bot-1201-54 Bark anatomy and cell size variation in Quercus faginea 1,2, 2 2 2 Teresa QUILHÓ *, Vicelina SOUSA , Fatima TAVARES , Helena PEREIRA 1 Centre of Forests and Forest Products, Tropical Research Institute, Tapada da Ajuda, 1347-017 Lisbon, Portugal 2 Centre of Forestry Research, School of Agronomy, Technical University of Lisbon, Tapada da Ajuda, 1349-017 Lisbon, Portugal Received: 30.01.2012 Accepted: 27.09.2012 Published Online: 15.05.2013 Printed: 30.05.2013 Abstract: The bark structure of Quercus faginea Lam. in trees 30–60 years old grown in Portugal is described. The rhytidome consists of 3–5 sequential periderms alternating with secondary phloem. The phellem is composed of 2–5 layers of cells with thin suberised walls and narrow (1–3 seriate) tangential band of lignified thick-walled cells. The phelloderm is thin (2–3 seriate). Secondary phloem is formed by a few tangential bands of fibres alternating with bands of sieve elements and axial parenchyma. Formation of conspicuous sclereids and the dilatation growth (proliferation and enlargement of parenchyma cells) affect the bark structure. Fused phloem rays give rise to broad rays. Crystals and druses were mostly seen in dilated axial parenchyma cells. Bark thickness, sieve tube element length, and secondary phloem fibre wall thickness decreased with tree height. The sieve tube width did not follow any regular trend. In general, the fibre length had a small increase toward breast height, followed by a decrease towards the top. Fibre width decreased with height in most of the cases, but in some trees a slight increase was noticed at the top. Key words: Quercus, bark, anatomy, fibre, sieve tube elements 1. Introduction Trockenbrodt, 1994; Quilhó et al., 2000; Jorge et al., 2000; The Portuguese oak (Quercus faginea Lam.) is a species Babu et al., 2010; Arıhan & Güvenç, 2011; Tavares et al., 2011). native to the western Iberian Peninsula and the North Only very limited information was published on the African countries of Morocco, Tunisia, and Algeria. The bark of the Quercus L. species. Bark anatomy of white species grows well in the temperature range of 15–25 °C in and northern red oak was first described by Chang the summer and from –4 to 8 °C in the winter, with annual (1954). Whitmore (1962) compared the bark structure precipitation between 350 and 2000 mm (Oliveira et al., and surface pattern of mature trees of Q. robur, Howard 2001). It is a medium-sized deciduous or semievergreen (1977) observed the bark structure of 11 Quercus species, tree growing to a height of 20 m and a diameter of 80 cm. Trockenbrodt (1991, 1994, 1995) studied the qualitative In Portugal, Quercus faginea coexists with other oak and quantitative anatomy of Q. robur bark, and Sen et al. species such as Quercus ilex L., Quercus suber L., Quercus (2011a, 2011b) described in detail the bark of Q. cerris L. pyrenaica Willd., and Quercus robur L. The distribution has var. cerris. One exception is the research effort and detailed become fragmented in the last centuries (Fabião & Silva, knowledge gathered on Q. suber bark, which is due to the 1996), and there are concerns on future area reduction economic importance of its cork component, as reviewed with warming and reduced rainfall trends, since drought by Pereira (2007). is the main limiting factor of sub-Mediterranean oaks The present study investigated the bark structure as (Cotillas et al., 2009), specifically for Q. faginea (Corcuera well as between- and within-tree size variation of the sieve et al., 2004; Montserrat-Marti, 2009). tube elements and secondary phloem fibres in Q. faginea. Studies on bark anatomy are scarce when compared The results are the first to be published on the bark of to previous studies on wood. However, knowledge on bark this species, thereby also adding new information on the structure and its variation within and between individuals structural characteristics of Quercus spp. barks. We also of a species, as well as age-related trends, are important to think that this knowledge, besides being necessary to assess bark’s diagnostic value and to determine its potential assess potential bark uses, has a role in the research on the uses (Roth, 1981). In this area, literature is also scarce (i.e. sustainability of Q. faginea in its indigenous regions. * Correspondence: [email protected] 561 QUILHÓ et al. / Turk J Bot 2. Materials and methods tangential diameter of parenchyma cells, sclereids, and The anatomical studies were made from the bark of 10 cluster of sclereids were also measured to estimate their randomly selected Quercus faginea Lam. trees, growing range of values. in pure stands in north-eastern Portugal, in the district of A light microscope Leica DM LA with camera Bragança (41°30′41″N, 7°01′06″W; altitude 554 m, 600– Nikon Microphot-FXA was used for light microscopic 800 mm annual rainfall, and 12–14 °C mean temperature). observations. Tree characteristics are shown in Table 1. The soils are Mean values for the tree were calculated as the mainly Orthic Dystric Leptosols on shale or granite rocks arithmetic mean of the 3 height levels in each tree. and Orthic Eutric Leptosols on basic rocks. The pH varies Analysis of variance (ANOVA) was applied to determine from acid to neutral, and the hard rock is found within 50 cm of the surface. if the differences of sieve tube elements and bark fibre size Cross-sectional 5-cm-thick discs were cut from each within and between trees were statistically significant at a tree along the stem at different height levels, and 2 cross- 0.05 confidence level. Statistical calculations were carried diameters were measured for bark thickness determination. out with IBM SPSS Statistics v.19 software. For qualitative anatomical characterisation, bark The descriptive terminology for bark structure follows samples were collected at breast height (bh), i.e. at Trockenbrodt (1990), Junikka (1994), and Richter et al. 1.30 m above ground. The samples were impregnated (1996). with DP1500 polyethylene glycol, and transverse and longitudinal microscopic sections of approximately 17 µm 3. Results and discussion thickness were prepared with a Leica SM 2400 microtome The bark of the Quercus faginea trees showed a scaly surface using Tesafilm 106/4106 adhesive (Quilhó et al., 1999). of grey-brown colour, with short but deep longitudinal The sections were stained with a double staining of furrows (Figure 1). The visual appearance was similar to chrysodine/astra blue. Sudan 4 was also used for selective other Quercus spp. (Whitmore, 1962; Howard, 1977). The staining of suberin. The stained sections were mounted on bark thickness ranged between 0.5 cm and 1.4 cm at bh Kaiser glycerine, and after 24 h drying, the lamellas were submerged into xylol for 30 min to remove the Tesafilm, (Table 1), corresponding to 10% of the total tree radius. were dehydrated on 96% and 100% alcohol, and were Table 2 presents the data for each tree showing that bark mounted on Eukitt. thickness tends to increase with age, i.e. higher values For quantitative studies, specimens were taken from occur in the base of the tree, agreeing with observations in bark samples collected at 3 tree height levels (base, bh, and other taxa (Trockenbrodt, 1994; Quilhó et al., 2000). There 75% of total height), and samples were taken sequentially was a significant difference between the bark thickness of from the cambium towards the periphery. Specimens were each tree at the 3 height levels, indicative of a significant macerated in a 1:1 solution of 30% H2O2 and CH3COOH axial variation of bark thickness in the tree. at 60 °C for 48 h and then stained with astra blue. Length, width, and cell wall thickness of 40 fibres and 40 sieve tube elements were measured on each specimen using a microscope and a semiautomatic image analyser. The number of measurements was previously calculated to give an accuracy of 95% at the 0.05 probability level. The Table 1. Characteristics of the Quercus faginea trees (mean and standard deviation, min. and max. of 10 trees). Mean ± std. dev. Min.–Max. Tree age (years) 40.4 ± 7.50 34–60 Total tree height (m) 10.5 ± 0.67 10–12 Height of living crown (m) 8.3 ± 1.34 6–11 Stem height (m) 2.2 ± 1.02 1–4 Diameter at bh (cm) 20.9 ± 4.20 16–29 Figure 1. Transverse section of Quercus faginea bark. Phm: secondary phloem; Pr: periderm, Rhy: rhytidome; arrow: clusters Bark thickness at bh (cm) 1.0 ± 0.29 1–1 of sclereids; Xm: xylem. Scale bar = 2 mm. 562 QUILHÓ et al. / Turk J Bot Table 2. Mean bark thickness of 10 Quercus faginea trees at 3 height levels (base, bh, and top). Bark thickness (cm) 1 2 3 4 5 6 7 8 9 10 Top 0.43 0.35 0.15 0.50 0.23 0.50 0.40 0.43 0.33 0.43 bh 1.18 1.03 0.50 0.75 0.63 1.20 1.23 1.38 0.85 1.00 Base 1.88 1.00 0.68 1.18 0.85 3.83 1.28 2.08 2.13 1.88 3.1. Bark structure the periderm was a consequence of the presence in its path The bark structure ofQ. faginea was identical in the 10 of sclereid clusters and fused rays (Figure 2). Q. faginea did trees. It was possible to distinguish macroscopically in not produce extensive phellem or cork layers, as is the case the bark cross-section the phloem, the periderm, and the in the discontinuous phellem of Q.
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