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Wood and Bark Anatomy of Alluaudia (Didiereaceae) from Madagascar

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Wood and Bark Anatomy of Alluaudia (Didiereaceae) from Madagascar IAWA Bulletin n.s., Vol. 1 (3),1980 133 WOOD AND BARK ANATOMY OF ALLUAUDIA (DIDIEREACEAE) FROM MADAGASCAR by R.W. den Outer and W. L. H. van Veenendaal Department of Botany, Agricultural University, Wageningen, The Netherlands Summary The anatomy of the secondary phloem and south of Tulear) is a shrub, 2 m heigh, a stem xylem of the tree Alluaudia ascendens Drake diameter of 4 cm and about 15 growth rings has been compared with that of Alluaudia near the stem base. Anatomical features were humbertii Choux, both belonging to the studied in transverse, radial and tangential endemic family Didiereaceae of Madagascar. sections and macerations. All sections were The differences between the two species are embedded in Kaiser's gelatin-glycerin (Johan­ small. In contrast with A. humbertii the cortex sen, 1940). Means and ranges of the length of of A. ascendens, persistent during the lifetime sieve-tube members, vessel members, parenchy­ of the tree, is thick and possesses a network of ma strands, fibre-sclereids, fibrous elements vascular bundles. These bundles are in con­ and the radial vessel diameter are based on at nection with the vertical system of the stem least twenty-five individual measurements. The via the horizontal ray system in non-conduc­ sieve-tube type and companion-cell type were ting and conducting secondary phloem and classified according to Zahur (1959). The air secondary xylem. The function of these dried material used for S.E.M. had clean cut bundles is discussed. surfaces, was sputter-coated with gold and observed in a Jeol scanning electron micro­ Introduction scope type U-3. The endemic family Didiereaceae from the dry area of south-west Madagascar, is a small Results one, composed of 4 genera with in all 11 or 12 species. The representatives are trees or shrubs Wood Anatomy somewhat cactus-like in habit with small simple The secondary xylem cylinder of the inves­ leaves or only thorns; they are clearly charac­ tigated Alluaudia ascendens has a diameter of terized by the architecture of the flowers. The about 10 cm. four genera of the family are Decaryia, Alluau­ Growth rings fairly distinct, (0.3-) 1.3 ( - diopsis, Didierea and Alluaudia. Information 2.3) mm (Fig. 5). Growth-ring boundaries are on their wood - and especially bark - anat­ marked by more and larger vessels in the early omy is very scarce. Samples of only two wood, while in the late wood libriform fibres Alluaudia species were at our disposal and used are flattened and the ray cells nearly always for this study. short in radial direction. The greyish-beige wood is rather often semi-ring-porous. Heart­ Materials and Methods wood is not recognizable. Bark and wood samples used are from the Vessels c. 40/mm2, solitary or in radial to Van Veenendaal/Den Outer collection, Mada­ tangential multiples or clusters, round to gascar (1978). The collection is housed at the oval or flattened where in contact with each Department of Botany, Agricultural University, other, radial diameter (40-)100( -200) 101m, Wageningen, The Netherlands. All the material tangential diameter (40-)80( -120) 101m. Walls studied is accompanied by herbarium vouchers. 1.5-2 101m thick. Vessel-memberlength (100-) The samples were collected from stems and im­ 250( -430) 101m. Perforations simple (sometimes mediately fixed in F.A.A. Alluaudia ascendens in pairs) in slightly oblique to horizontal Drake (collection number V. and O. 1129, 10 end walls. Inter-vessel pits alternate, 6-7 km east of Tsihombe) is a tree of 8 m heigh, 101m, sometimes with coalescent apertures. with a dbh of II cm and about 35 growth rings Vessel-ray and vessel-parenchyma pits half­ at the stem base. Alluaudia humbertii Choux bordered to almost simple, 10-25 101m. Vessels (collection number V. and o. 1068, 18 km usually in contact with parenchyma. Downloaded from Brill.com09/25/2021 04:54:15PM via free access 134 IAWA Bulletin n.s., Vol. 1(3),1980 Fig. 1-4. Alluaudia ascendens Drake. Scanning electron micrographs of the secondary xylem. -- I. Transverse and tangential surface, showing a large wood ray with tracheal elements in its centre, x 145. -- 2. Radial surface. Tracheal elements of the vertical system bend towards and make contact with tracheal elements of the wood ray. Note the simple crystals, x ISO. -- 3. Transverse surface, showing tracheal elements in the wood ray. Note the simple perforation in the centre, x 510. -- 4. Transverse surface, showing spiral thickenings in tracheal wood-ray elements and parenchyma cells, x 5240. Downloaded from Brill.com09/25/2021 04:54:15PM via free access IAWA Bulletin n.s., Vol. 1 (3), 1980 135 Librifonn fibres (600-)880( -1190) Mm phloem immediately outside the cambial zone long, walls 3 Mm thick (in the late wood (200 Mm), the non-conducting phloem (620 somewhat thicker) with many simple pits with Mm), a cortex (c. II mm) and finally the peri­ slit-like inner apertures mainly confined to the derm (c. 200 Mm). radial walls often in two vertical rows, non­ The axial system on cross section of the septate. Fibre-tracheids and vascular tracheids conducting secondary phloem (Figs. 9 & 10), frequent near the vessels, c. 470 Mm long, dia­ is composed nearly half of sieve tubes and meter 25 Mm. companion cells and about half of parenchyma ParenchYmil paratracheal (usually forming a cells. Fibres (fibre-sc1ereids) are absent or complete sheath of one cell wide around the scarcely present in the transitional zone to the vessels), apotracheally diffuse and marginal non-conducting phloem. (initial). Strands of 2-3 cells, c. 500 Mm Sieve-tubes scattered or in groups of 2 or long. 3 (bast type s or g; Den Outer & Fundter, Rays (4-)6(-9)jmm, 2-3-seriate, usually 1976), diameter c. 22 Mm. Sieve-tube member with short uniseriate tails, composed of erect type II (Zahur, 1959) length (165-)335(-430) (lg 120, rd 80, tg 45 Mm), square (lg 80, rd 80, Mm. Sieve plates compound (c. 4 sieve areas), tg 45 Mm) and a few procumbent cells (lg 70, oblique to horizontal, many starch-like grains rd 120, tg 45 Mm). Width c. 90 Mm, height which do not colour purple when treated with 1100(-2000) Mm. Perforated ray cells rather iodine near the sieve plates (Fig. 11). Sieve often present, perforations up to 50 Mm. areas in radial walls, indistinct, scarce, horizon­ Some rays are much wider, c. 280 Mm (Figs 1 & tal diameter 6 Mm. 7) with in their centre vessels with spiral Companion cells type C (Zahur, 1959), thickenings, vascular tracheids and tracheids 3( -2) cells per strand, oval on cross section, (Figs. 3, 4 & 8) not accompanied by phloem diameter 7-8 Mm. Strands usually situated elements. The horizontally stretched tracheal along one of the radial walls of the sieve-tube elements (length (100-)150(-230) Mm, member. diameter 10-15 Mm) within the rays bend at a ParenchYmil cells round to oval on cross sec­ certain point into a vertical direction, leave the tion or flattened in radial direction, radial ray and become part of the axial system (Figs. and tangential diameter 22-45 Mm, red-brown 2 & 6). to dark brown contents absent (in contrast Crystals in ray parenchyma cells rhomboidal; with the non-conducting phloem). Strands of 2 druses less frequent. cells, length (390-)440( -480) Mm. Rays uniseriate and 3( -2)-seriate with short The secondary xylem of the investigated uniseriate tails, composed of erect and square Alluaudia humbertii differs slightly from that cells (lg 65, rd 65, tg 50 Mm) and a few pro­ of A. ascendens. The following differences cumbent ones. Cell contents red-brown sub­ were observed. stances or druses. Some of the multiseriate rays Growth rings more distinct, (0.2-)0.4( -0.5) are more than 3 cells wide and possess radial mm, diffuse vessel arrangement. All axial ele­ vascular bundles (Figs. 14 & 16). ments are somewhat shorter. Vessels c. 5 2jmm 2, The non-conducting phloem consists of dila­ radial and tangential diameter one fourth tating phloem rays of the TWa-type alter­ smaller, perforations more often in pairs. nated with wedge-shaped areas composed of Walls 2.5-3.5 Mm. Inter-vessel pits more often elements of the axial system. The sieve tubes opposite. Vessels usually in contact with li­ are crushed in these areas. The parenchyma briform fibres. Libriform fibres with simple cells sometimes have red-brown contents. Many pits in one vertical row, walls 5 Mm thick. fibre-sc1ereids are formed, length (350-)410- ParenchYmil mainly diffuse and initial. Rays (-470) Mm, walls thick (7 Mm) in which many 5 jmm, usually 4-seriate, width c. 180 Mm, simple pits with slit-like vertical inner aper­ height c. 1300 Mm, composed of erect (lg 100, tures. rd 45, tg 45 Mm) and square cells (cubical, The cortex is almost entirely composed of 45 Mm wide). Procumbent cells, perforated ray parenchyma cells. Towards the periderm rela­ cells and wide rays with tracheal elements in tively thin-walled sc1ereids are present. Paren­ the centre, absent. Crystals present as sand, chyma cells tangentially stretched (tg 165 Mm) seldom druses, in the ray parenchyma cells. (strong dilatation). Contents rather frequently a large druse or red-brown substances especially Bark Anatomy near the periderm, sometimes dark brown to The bark of the investigated Alluaudia black I!;ranular masses. Some parenchyma cells ascendens is about 12 mm thick. It can be are much larger (diameter c. 240 Mm), relative­ divided into four zones, i.e. the conducting ly thick-walled without any visible contents. Downloaded from Brill.com09/25/2021 04:54:15PM via free access 136 IAWA Bulletin n.s., Vol.
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