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Heidelberglaan 2, Tafrique Anisophyllus Ray' Xylem 113 The occurrence of axial latex tubes in the secondary xylem of some species of Artocarpus J. R. & G. Forster (Moraceae) by S.M.C. Topperand J. Koek-Noorman Institute of Systematic Botany, University of Utrecht, Heidelberglaan 2, Utrecht, The Netherlands Summary A. anisophyllus Miq.: Sabah, Uw 24213, In a number of species ofArtocarpus J.R. & SAN 50567; Indonesia, Uw 24607 (RTIw, IND G. Forster (Moraceae) laticifers were not only coll. 2307), BO 1778; S.E. Asia, Uw 24308, observed as radial tubes, enclosed in the ray BO Pfeiffer E 952. — A. blancoi (Elmer) Merr.: tissue, but also as axial tubes, enclosed in Philippines, Uw 24214, Tw Rojo 262*. — A. communis L the fibre tissue. Both types of latex tubes are Forst.: New Guinea, Uw 24325, L BW Uw connected and considered as one branched lati- BW 1465 (sterile); Uw 24603, 4219; L BW Uw L BW ciferous system. A detailed description of the 24600, 1269; 24602, 2904; individual latex tubes and the whole system, Uw 24326, L BW 2541; Samoa, Uw 16632,US based on light microscopic and scanning elec- coll. L.S. Dutton nr. 8;Philippines, Uw 24310, Pfeiffer TS*. — tron microscopic observation, is presented. BO 3113 A. elasticus Reinw. ex Blume: Java, Uw 24421, BO Koorders coll. Introduction 1026c/2203lb; Uw 24422, BO Koorders coll. The present investigation is part of a de- 1621m*;Sabah, Uw 24215, SAN 18656. - A. tailed, comparative wood anatomical study of fretessii T. & B. in Hassk.: New Guinea, Uw the Moraceae. 18112, Lw BW9871*;Uw 24318, Dw Fokkin- Until now latex tubes in the secondary ga 2175*; Uw 24316, L BW 2133. -A. integer xylem of Moraceae and in general have been (Thunb.) Merr.: New Guinea, Uw 24319, L BW observed only in the rays. Consequently a latex 5659; Philippines,Uw24323,BO Pfeiffer 385*. defined tube in the secondary xylem has been — A. kemando Miq.: Sumatra, Uw 24324, L En- of the by the Committee of Nomenclature dert & v.d. Zwaan nr. 90, R.R.I. 755; Indone- IAWA (1964) as ‘a laticifer enclosed in a ray’. sia, Uw 24605 (RTIw IND coll. 3590)*;Malaya, In our search of the literature we found two Uw 24608, FHOw 4536, Desch FMS 28853*. considered them exceptions but both authors — A. lakoocha Roxb.: E. Pakistan, Uw 18033, as an exception of the rule: Tippo (1938) MADw 24487. — A. lanceifolius Roxb.: Moluc- remarks that of the radial latex tubes of Uw L Vriese some cas, 24609, De & Teysmann 20; Craterogyne kameruniana (Engl.) Lanj. (= Sabah, Uw 24216, SAN 59079; Indonesia, Uw Dorstenia, Berg, 1978), are ‘outside of the ray’ 24604 (RTIw IND coll. 2200), BO 1682; Uw and Vreede (1949) gives anexample of a radial 24606 (RTIw IND coll. 4027)*; S.E. Asia, Uw not latex tube that ‘does lie in a ray’. As far as 24328, Dw Pfeiffer 6891*. - A. nitidus Tree.: known to us, axial latex tubes have never been Sabah, Uw 24217, SAN 25510. — A. sepicanus reported for the secondary xylem. Diels: New Guinea, Uw 24329, Dw Fokkinga 7822*. — A. sericicarpus Jarrett: Sabah, Uw Materials 24218, SAN 50586. — A. tamaran Jarrett: Sa- Only dried material was used for this inves- bah, Uw 24219, SAN 25581. — A. teysmannii tigation. The material was partly present in the Miq.: New Guinea, Uw 24330, L BW 1204. Utrecht wood collection and was partly obtain- The available slides in our institute and in ed by the kind cooperation of the curators of the Rijksherbarium Leiden have been used in the wood collections of the search of axial latex tubes in other of Rijksherbarium genera Leiden, the Royal Tropical Institute Amster- Moraceae. The numbers between brackets indi- dam, TNO Houtinstituut Delft and the Musee cate the number of species studied: Antiaris Royal de TAfrique Centrale Tervuren. The (10); Bagassa (3); Batocarpus (4); Bosqueiopsis genus Artocarpus is represented by the follow- (1); Brosimum (12); Broussonetia (3); Castilla ing species. An asterisk indicates the absence or (6); Chlorophora(7); Clarisia (3); Cudrania (3); uncertainty about the existence of herbarium Dorstenia (1); Ficus (5); Helianthostylis (2); vouchers. The abbreviations used are according Helicostylis (9); Maclura (1); Maquira (7); to the Index Herbariorum (Holmgren&Keuken, Morus (3); Naucleopsis (7); Olmedia (2); Parar- 1974) and the Index Xylariorum (Stem, 1978). tocarpus (2); Perebea (3); Plecospermum (1): 114 lAWA Bulletin n.s., Vol. 1 (3), 1980 Prainea (2); Pseudolmedia (5); Sorocea (3): Frequency of the occurrence of the latex Sparattosyce (I); Streblus (3); Treculia (I); tubes — The frequency of occurrence of the Trilepisium (2); Trophis (2); and Trymatococ- axial latex tubes is rather low. Although exact cus (3). figures are not available a rough estimate is here given: in a cross section of about one square centimeter one may expect 0-6 latex Methods tubes. Radial latex tubes occur generally more Sudan III was used for staining latex as frequently but sometimes they are as rare as recommended by Vreede (1949). The scanning axial latex tubes or not present at all (Table 1). electron microscope, used in this investigation All wood samples with axial latex tubes also Stereoscan S 604. Photo- was a Cambridge showed radial latex tubes. at made 7.5 Kvolt. Microtome The the - graphs were contents of latex tubes The con- sections of about 20 microns thick first tents often were are yellowish brown, granular, re- with the aid of the micro- the contents photographed light sembling dark cell of many paren- Afterwards scope. they were air dried, sputter chyma cells. Upon staining with Sudan III the coated with gold and studied with the SEM. latex in the latex tubes turns into orange red, This enabled us to compare light micrographs and the contents of the parenchyma cells be- and SEM observations, and thus facilitated the come purplish red. detection of the axial latex tubes the among fibres with the scanning electron microscope. Table 1. Presence (+) or absence (—) of axial (a.l.t.) and radial latex tubes (r.l.t.) in the Arto- Results carpus material studied. The occurrence of axial and radial latex tubes — In agreement with data from the litera- Sample a.l.t. r.l.t. ture (Solereder, 1885; Janssonius, 1934 as A. anisophyllus Uw 24213 + + ‘radial laufende Elemente’; Metcalfe & Chalk, yy Uw 24607 + + »» 1950;Brazier & Franklin, 1961;Sharma, 1962; Uw 24308 — + Burgess, 1966;Kribs, 1968; Purkayastha et al., A. communis Uw 24325 + + »» Uw 24603 1976) radial latex tubes were observed in a + + number of yy Uw 24310 + + species of the large genus Artocar- yy of 52 Uw 24600 — — pus, consisting species (Jarrett, 1977). In the secondary xylem of seven out of the four- »» Uw 24602 — — teen »» Uw 24326 — — species (see Table 1) latex tubes occur »» not Uw 16632 — — only in the rays, as they usually do, but also in axial direction in the fibre tissue. A. integer Uw 24319 + + Not all specimens of these seven species show- »» Uw 24323 — — ed axial latex tubes. Neither have axial latex A. kemando Uw 24324 + + tubes been »» Uw 24605 observed until now in the other + + » listed species studied of Artocarpus, nor in Uw 24608 — + other A. lanceifolius Uw 24609 any moraceous genus. + + Radial latex tubes on the other hand have »» Uw 24606 + + been recorded for specimens of all Artocarpus yy Uw 24604 + + species studied. yy Uw 24328 + + The position of the latex tubes in the sec- yy Uw 24216 — + ondary xylem — In Artocarpus radial latex A. sericicarpus Uw 24218 + + tubes are always surrounded by ray parenchy- A. teysmannii Uw 24330 + + ma. The axial latex tubes lay scattered in the A. blancoi Uw 24214 — + fibre tissue and are consequently surrounded A. elasticus Uw 24421 — — by fibres (Fig. 1). They only make lateral con- Uw 24422 — — » with the their Uw 24215 — tact rays in course through the + fibres. The lumina of the axial latex tubes are A. fretessii Uw 18112 — + »» only slightly larger than the lumina of the Uw 24318 — + »» largest fibres and of the parenchyma cells Uw 24316 — + which A. lakoocha 18033 — + makes them barely recognizable in cross Uw sections (Fig. 1). However, at high magnifica- A. nitidus Uw 24217 — + tions, the differences in size, wall thickness and A. sepicanus Uw 24329 — + cell contents are quite striking. A. tamaran Uw 24219 — + lAWA Bulletin n.s., Vol. 1 (3), 1980 115 The constitution of the wall of the latex Discussion tubes — The walls of the laticifers are diffi- Articulated or nonarticulated laticifers — cult to observe when they are filled with latex. Already David (1872) distinguished between However, often the tubes are partly empty or articulated (Milchsaftgefasse) and nonarticu- the latex is removed during sectioning. Both lated (Milchsaftschlauche) laticifers. The tubu- with the light microscope and the SEM the lar laticifers of the Moraceae are commonly most existence of a thin wall is obvious in considered as nonarticulated laticifers, compa- those places where it is locally ruptured during rable with those in the Euphorbiaceae, Asclepi- sectioning (Fig. 2 & 3), or as happened once adaceae, Apocynaceae and Urticaceae (David, during the observation with the SEM, where 1872; Schaffstein, 1932; Vreede, 1949). two blind pits were blown up as a result of However, Dippel (1865) and Milanez (1954) local gas production (Fig. 4). The walls of the defended the ‘articulated’ point of view, based to laticifers, although distinct, are extremely thin, upon studies of Ficus. We are inclined con- less than 0.5 micron thick (Fig. 2 & 3). Pits sider the latex tubes of Artocarpus as ‘non- were found neither in in axial articulated’ because walls radial, nor no cross or per- latex tubes. Radial latex tubes show often forated cross walls could be found. Some cau- to blind pits contiguous parenchyma cells (Fig. tion remains necessary as we observed with the In macerations of latex tubes much 4).
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