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Wound-Induced Periderm Tubes in the Bark of Melia Azedarach, Ficus

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Wound-Induced Periderm Tubes in the Bark of Melia Azedarach, Ficus IAWA Bulletin n.s., Vol. 12 (1),1991: 62-66 WOUND-INDUCED PERIDERM TUBFS IN THE BARK OF MELIA AZEDARACH, FICUS SYCOMORUS AND PLATANUS ACERIFOLIA by Simcha Lev-Yadun and Roni Aloni Department of Botany, The George S. Wise Faculty of Life Sciences, Tel Aviv University, Tel Aviv 69978, Israel Summary Tubes of periderm were induced experi- lating mechanism for the formation of these mentally by wounding sterns of Melia aze- periderm tubes was not suggested (Wisniew- darach L., Ficus sycomorus L. and Platanus ski et al. 1984). Periderm tubes were also acerifolia Willd. This wounding stimulated found in the horned oak galls induced on the formation of periderm tubes around those Quercus palustris by Callirhytis corniqera. In strands of phloem fibres which were exposed this case the periderm tubes were not associ- to the outside atmosphere. The differentiation ated with fibres. Instead, they encircled the of periderm tubes inside the bark is a defence insect ovum and compartmentalised it from reaction by which the wound-exposed fibres the other living cells in the tissue (Taft & Bis- are isolated from the living cells of the bark. sing 1988). In several Pinus species, tubes of The regulating factors of wound-induced periderm were formed around the wounds periderm tubes are probably ethylene and at- induced by pi ne bast scale (Matsucoccus mospheric air. iosephi) infestation and by needle punctures Key words: Ficus sycomorus, Melia aze- (Liphschitz & Mendel 1989). In intact sterns darach, periderm differentiation, Platanus of Vitis vinijera, periderm tubes naturally acerifolia, wound response. occurred around secondary phloem fibre strands which were partly exposed to air Introduction (Aloni & Peterson 1991). The phellogen, which is the meristematic Here we show that tubes of periderm can cell layer of the periderm, usually produces be induced experimentally around phloem strips, sheets or scales of phellern. Phellogen fibres exposed to air by wounding. We in- initiation generally occurs in the older parts duced periderm tubes in three tree species of sterns and roots and serves as a secondary and suggest a controlling mechanism for their protective tissue. In addition, phellogen is differentiation. formed following wounding (Fahn 1982). Infrequently, periderms are formed in the shape of tubes inside the bark. Early descrip- Materials and Methods tions of this type of periderm used the term Twenty branches of each of the following 'Doppelperiderm' (double-periderm). In La- species: Melia azedarach, Ficus sycomorus burnum adami, this doubled-periderm had a and Platanus acerifolia, grown near Tel Aviv central core of collenchyma and was oval in a University, were wounded during the spring transverse section (Buder 1911). Tubes of of 1989 wilh a sharp razor blade. The wounds periderm were also found in several cases of reached the cambium and were of various tissue patho10gies. In wounded peach trees, shapes, forming either narrow vertical bark primary phloem fibres 3-5 rum from the strips, or vertical, horizontal and oblique wound margin were circumvented by finger- scratches. The wounded organs as weil as like projections of necrophylactic periderm. untreated control branches were sampled 2- However, an explanation conceming a regu- 12 months after wounding. They were then Downloaded from Brill.com10/02/2021 04:29:33AM via free access Lev-Yadun & Aloni - Wound-induced peridenn tubes 63 sectioned with a sliding microtome (trans- encircled by a single large peridenn tube verse and longitudinal sections), stained with (Fig. 4). In radial longitudinal sections, the safranin and fast green, and mounted in longitudinal patterns of the peridenn tubes Canada balsam (Jen sen 1962). The sections were observed around secondary phloem were studied with a Leitz Dialux-20 micro- fibre strands which were exposed to air scope, equipped with a Nicon F-3 camera. (Figs. 5 & 6). In Ficus sycomorus and Platanus acerifo- lia, a few wound-indueed tubes of peridenn Results differentiated around the strands of phloem In each of the three species studied, peri- fibres which were exposed to air by a cut derm tubes were fonned around strands of (Figs. 7 & 8). phloem fibres which had been exposed to air by the wounding process. Tubes of peri- denn were not observed in the intact control Discussion branches of these three species. In Melia aze- The present study shows that the differen- darach, numerous periderm tubes were in- tiation of wound-induced tubes of peridenn duced by wounding (Figs. 1 & 2). Near the following wounding is a general phenom- surfaee of the wound, fonns intermediate be- enon oecurring in woody plants with fibre tween a periderm tube and a peridenn sheet strands in their bark. In eaeh of the three spe- were often detected (Fig. 1). In narrow brid- eies studied, the wound-indueed periderm ges of bark, whieh enhanced the wound ef- tubes were assoeiated with those phloem feet, the periderm tubes were relatively thick fibres exposed to the outside surrounding at- (Figs. 1-3) and had a maximum of 12 suber- mosphere. In previous reports of periderm ised celllayers in the wall of a tube (Fig. 3). tubes, these structures were usually associated Most of the exposed fibre strands were sur- either with fibres in Vitis vinifera (Aloni & rounded by a single periderm tube. How- Peterson 1990) and Prunus persica (Wisniew- ever, some of the tubes united tangentially ski et al. 1984), or with wounding in Quer- forming a large eontinuous periderm tube cus palustris (Taft & Bissing 1988) and Pinus which encircled more than one group of fi- speeies (Liphschitz & MendelI989). bres (Figs. 1-3). Oecasionally, a few radial In our study, as weil as in the case of the strands of seeondary phloem fibres were naturally occurring peridenn tubes in Vitis Legends for Figures 1-8: ---7 Figs. 1-4. Melia azedarach; bark; TS. - 1: Wound-induced peridenn tubes (arrows) around strands of secondary phloem fibres (F). Note intennediate patterns (arrowheads) between peridenn tubes and a peridenn sheet at the surfaee of the wound (W). - 2: Numerous wound- induced peridenn tubes indueed in a narrow bark strip of a partially girdled stern. Note tangen- tially enlarged tubes with two fibre strands per peridenn tube (arrows). - 3: Close-up view of radially conneeted wound-indueed peridenn tubes with many phellern layers (arrows) around strands of fibres. - 4: A large wound-indueed peridenn tube (arrows) engulfs three strands of phloem fibres immediately beside a wound. Seale bar :, 250 ~m. ---7 ---7 Figs. 5 & 6. Melia azedarach; bark; RLS, showing the same wound-indueed peridenn tube (arrows) fonned around a strand of seeondary phloem fibres (F) exposed to air by a horizontal wound (W). - Fig. 7. Ficus sycomorus; bark; TS, showing a wound-induced eork tube (ar- row) fonned around phloem fibres. - Fig. 8. Platanus acerifolia; bark; TS, showing two adja- cent wound-indueed periderm tubes (arrows) surrounding phloem fibres. Seale bar = 250 ~m. Downloaded from Brill.com10/02/2021 04:29:33AM via free access 64 IAWA Bulletin n.s., VoI. 12 (1),1991 Downloaded from Brill.com10/02/2021 04:29:33AM via free access Lev-Yadun & Aloni - Wound-inducedperiderm tubes 65 Downloaded from Brill.com10/02/2021 04:29:33AM via free access 66 IAWA Bulletin n. s., Vo!. 12 (1), 1991 vinifera (Aloni & Peterson 1990), atrnospheric Buder, J. 1911. Studien an Laburnum Adami. air penetrated into the bark tissues along the H. Allgemeine anatomische Analyse des exposed fibre strands subsequently drying Mischlings und seiner Stammpflanzen. them. It is likely that air penetration was also Zeitschr. f. Induktive Abstammungs- und involved in the formation of periderm tubes Vererbungslehre 5: 209-284. in infected Prunus persica tissues (Wisniew- Fahn, A. 1982. Plant anatomy. 3rd ed. Perga- ski et al. 1984) and in Quercus palustris (Taft rnon Press, Oxford. & Bissing 1988) where air entered the tissue Jensen, W.A. 1962. Botanical histochemis- through ovipositor tracks. Aeration or expo- try, principles and practice. Freeman & sure to oxygen is known to induce the initi- Co, San Francisco. ation of phellogen and to promote its activity Lev-Yadun, S. & R. Aloni. 1990. Polar pat- (Priestley & Woffenden 1922; Liphschitz & terns of periderm ontogeny, their relation- Waisel 1970). Thus, formation of tubes of ship to leaves and buds, and the control periderm is probably simply a specific case of cork formation. IA W ABui!. n. s. 11: of the general effect of aeration on periderm 289-300. formation. Liphschitz, N. & Z. Mende!. 1989. Interac- The wound effect is another factor which tion between hosts and non-hosts ofPinus should be considered in our experimental sys- spp. and Matsucoccus josephi: anatomical tems. Periderm is usually induced around responses of stern to infestation. New wounds (Fahn 1982) compartrnentalising dam- Phyto!. 113: 135-142. aged tissues from unaffected ones (Biggs Liphschitz, N. & Y. Waise!. 1970. Phellogen 1985). Plant tissues synthesise ethylene in initiation in the sterns ofEucalyptus camal- response to wounding (Yang & Hoffman dulensis Dehnh. Aust. J. Bot. 18: 185-189. 1984), ethylene being the major hormone Priestley, J.H. & L.M. Woffenden. 1922. which induces phellogen initiation and pro- Physiological studies in plant anatomy V. motes periderm formation (Lev-Yadun & Causal factors in cork formation. New Aloni 1990). Therefore, the two regulating Phyto!. 21: 252-268. factors which induced the formation of tubes Taft, J.B. & D.R. Bissing. 1988. Develop- of periderm in our experiments were proba- mental anatomy of the horned oak gaU in- bly wound-induced ethylene and atmospheric duced by Callirhytis cornigera on Quercus air. palustris (pin oak). Amer. J. Bot. 75: 26- 36. References Wisniewski, M., A. L. Bogle & C. L. Wilson. Aloni, R. & C.A. Peterson. 1991. Naturally 1984. Histopathology of canker develop- occurring periderm tubes around second- ment on peach trees after inoculation with ary phloem fibres in the bark of Vitis vini- Cytospora leucostoma.
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