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Wound Reactions in Bamboo Culms and Rhizomes*

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Wound Reactions in Bamboo Culms and Rhizomes* Journal of Tropical Forest Science 7 (1997 39 9(3)- 9 ) 37 : WOUND REACTION BAMBOSN I O CULM RHIZOMESD SAN * G. Weine Lies. W er& Universitat Hamburg, Leuschnerstrasse 91, 21031 Hamburg, Germany Received January 1996_________________ _______________ WEINER, G. & LIESE, W. 1997. Wound reactions in bamboo culms and rhizomes. Bamboo culms and rhizomes respond to wounds in order to protect the surrounding tissues against damaging influences throug woune hth d surfaces defence .Th e arsenal consist numbea f so f cellulao r r reactions suc closurs ha f sieveo e tube callosey b s , formatio slimf no tylosesd ean , phenolics, suberised wall layers, wall lignificatiod nan also septa developmen fibresn i t . These response similae sar thoso rt hardwoodsn ei , but due to lack of a secondary meristem, neither a barrier zone nor a callus for wound closine developedb n gca . Ther alss ei o accumulatio d mobilisationan f starcno h around the wound, development of additional lamellae of the cell wall in parenchyma formatiocelle fibresd th san d an suberi,a f no n laye vasculan ri r parenchyma cellse .Th spatial expansion of the wound reaction shows a distinct lateral boundary, but axially, it fades out gradually. Key words: Bambo owoun- d reaction sanatom- ycul- m rhizom- e WEINER, G. & LIESE, W. 1997. Reaksi kecederaan dalam rumpun dan rizom buluh. Rumpun buluh dan rizom buluh bertindakbalas terhadap kecederaan untuk melindungi tisu-tis i sekelilingnyud a daripada pengaruh yang merosakkan melalui permukaan yang cedera. Ketahanan arsenal mengandungi sebilangan reaksi sel seperti penutupan tiub tapis oleh kalusa, pembentukan lendir dan tilosis, fenol, lapisan dinding bergabus, penligninan dinding dan juga perkembangan septa dalam gentian. Tindakbala sami in s a dengan yang terdapat dalam kayu keras, tetapi akibat kekurangan meristem sekunder, kedua-dua zon penghalang dan kalus bagi menutup kecederaan tidakdapatdikembangkan. Terdapatpengumpulan dan pergerakankanji i sekelilind g tempat yang cedera, perkembangan lamela tambahan dindin padl gse a sel parenkim n gentianda a n pembentukada , n lapisan gabus dala l parenkimmse a vaskular. Pengembangan spatial bagi reaksi kecederaan menunjukkan batas sisi yang ketara, tetap beransur-ansua i r hilang secara paksi. Introduction Plant organs have a natural protection against injury, like the bark for trees and the cortex for monocotyledons. If there is damage of the phloem and xylem, protective reactions restrict danger from invadin micro-organismsd an r gai dicotyledonr .Fo - ous trees, defence mechanisms such as compartmentalisation and callus formation have been intensively investigate recenn i d t years (e.g. Shig Maro& x 1977, Bauch et al. 1980, Shigo 1984,1994, Liese & Dujesiefken 1989, 1996, Bonsen & Kucera 1990, Schmitt & Liese 1990, Blanchette & Biggs 1992). The understanding of wound response was much enhanced by the CODIT Concept: Compartmentalisa- tion of Decay in Trees (Shigo & Marx 1977). *Dedicated to Professor Dr. Joseph Bauch on the occasion of his 60th birthday. 379 0 38 Journal of Tropical Forest Science 7 (199739 - 9(3) 9 ) 37 : Corresponding information about the reactions in monocotyledons is more meagre Hadidl E . i (1969) investigate woune dth d healing proces somn si e flower- plantsg in , including monocotyledons. Wound responses were reportee th n di rhizome of bananas by Van der Molen etai (1977) and in the stem of the royal palm (Roystonea regia) by Weiner and Liese (1995), but nothing is known about the defence mechanism f bambooso growts It . completes hi d withi monthsw fe na t bu , transpore th t systefunctioo t s decada m r ha longernfo d ean . Althoug hbamboa o culm appears rather resistant against physica hards l it damag o t , siliceoue edu s cortex, insect penetratn sca cule eth m wal e breelacuno t lth n di a (Singh 1990, Kovacs & Azarae 1995). Moreover, a bamboo plant suffers seriously during the planting process. The common vegetative propagation by culm, branch or rhizome cuttings leads to considerable injuries. Rhizomes of leptomorph species are cut to limit growth. The upper crown of newly planted bamboos is often cut off to reduce respiration bamboe par p th Chin n f to I . o t e oath culm removes i s routina s da e operation to avoid snow breakage and to utilise additional material. Also injuries, as during harvesting, cannot be excluded. In each case, the transport of water and sap in the vascular bundles is interrupted. Bamboo as a monocotyledon cannot develo w cellulapne r pathway callua r r wouno s fo s d closura laco t f ko e edu secondary meristem. Nevertheless, the damaging influence of the invading air must be efficiently blocked off to protect the surrounding tissue. Even branches must have such defence mechanism dyina seao f st of lg part fro remainine mth g culm. reactione Th bambof so o were therefore investigate artifician do woundt se l t sa e cul d rhizomth man numbea f o e f specieso r d als t brokean oa , n branches. Material and methods For this study , 380 bamboo samples belonging to ll genera were examined (Table 1). The bamboo culms were of different ages and from two different climatic locations - wer .t wit Woundmm ese drillinha 4 2- f o sg machine th n i e middle of an internode, at a basal node and in the rhizome at different times of yeae th r (Figure evaluatioTh . e1) responsf no e between woundin samplind gan g varied between one day and 360 days. Table 1 and Figure 1 present details of the wounding procedure. As a control, an unwounded culm from each species was woundee prepareth s sama e y th den d wa i material . For light microscopy (LM) all wounded tissues with an adjoining part above belod an w were cut, fixe storeformaldehydd % d4 an n di . Longitudina°C 4 t ea l sections aroun wounda d wer witt ecu slidinh a g microtome generar Fo . l observa- tions, the sections were double-stained with acridine/chrysoidine red and astra blue. For fluorochroming, an aqueous 0.1 % solution of acridine orange was used. Suberin was tested with sudan III+IV, callose "plugs" with aniline blue and starch with Lugol's solution. Ruthenium red reacted with pectinophile substances, iron chloride (FeCrlll) with phenolic substances. Further, phloroglucin/HCL indi- cated lignified cell walls. All sections were embedded in glycerine and examined wit Olympun ha microscop2 sBH e equipped with fluorescence. Tabl . Locatione1 preparatiod san investigatee th f no d material g rf Bamboo species Location Plan e ag t Wound area Seaso samplinf no g Observation Method of — (y) after wounding investigation ^ (days) _! cV Phyllostachys heteroclada 3, 7, 14 £. Pleioblastus maculalus Nanjing/China 3-4 15.1 Oct. - Dec. 1992 21,28 LM, longitudina1 2 l Pseudosasa amabilis 40,51 s. CX3 -Juny Ma e Semiarundinaria1993 . I . 9 d an fasluosa . 4 2 1- Bot. Garden H H , 1, 3, 7, 10, 14 , longitudinaLM j l Phyllostachys uiridiglaucescens 21,29 S 1 to Arundinaria fastuosa s Chusquea quielea 00 Fargesia muriela o Phyllostachys decora O3 Phyllostachys flexuosa 5 Phyllostachys nigra Q Phyllostachys propinqua Bot. Garden, HH 2-3 15. I. June 1993 7, 14 LM, transverse [g Phyllostachys quilio C5 Phyllostachys rubromarginata Sasa palmata Semiarundinaria fastuosa Sinarundinaria nilida Thamnocalamus spalhaceus Chusquea quielea Bot. Garden, HH Culm from 2. N. and 15. 1. Aug. - Sept. 1993 1,3,7, 10, LM, longitudinal Sasa palmata 1993 4. N. and 9. I. 14,21,28 Semiarundinaria fastuosa 2. N. and 9. I. Dendrocalamus spp. 4 d an 2 UppeGreenhouse . I . 3 r H JunBF , e 1993 7, 14 Sinarundinaria nilida BFH 2-3 15.1. July 1993 1, 3, 7, 10, LM, longitudinal 14,21,28 (continued) °° Tabl (continuede1 ) Chusquea t/uielta 15. I. Phyllostachys viridiglaucescens 3 2- Bot. Garden H H , 9. I . Jun -Jun3 e9 4 e9 360 LM, longitudinal Semiarundinariafasluusa 9. I. Thamnocalamus spathaceus 1 Bot. Garden H H , -Augy Ma . 1994 . I . 9 Eac dayh7 s LM, longitudinal Phyllostachys quilio up to 90 days Phylloslachys viridiglaucescens Bot. Garden, HH 199y 4Ma 3- 4. N . 9 d an . N . 4 7, 14,21 LM, longitudinal Phyllaslachys pubescens Nanjing/China Rhizome ' Aug. - Sept. 94 1,3, 7, 14 LM, transverse 21,28,42 1 of Sinarundinaria nitida1 BFH H H , 9. + 15. 1. April - May 1994 3,7, 14, TEM, transverse Tropical 21,28 longitudinad an l Phylloslachys viridiglaucescens Bot. Garden, 1 - 2 9.1. Dec. 94 -Jan. 95 7, 14,21,28 TEM, transverse GreenhouseH H , 9. I. Jan. - Feb. 95 TEM, transverse Fores BFH = Bundesforschungsanstalt fur Forst- u. Holzwirtschaft, HH = Hamburg, I. = internode, N. = node, LM = light microscope, transmissio= M TE n electron microscope. oo -4 I(DO Journal of Tropical Forest Science 9(3): 379-397 (1997) 383 Figure 1. Wounding pattern in a bamboo culm at an internode and node r transmissioFo n electron microscopy (TEM), small sample) s (lx'mm 4 1X aroun woune dth d area were processed. Samples were immerse day3 r n i sdfo Karnovsky's fixing solution (Karnovsky 1965), washed in 0.1M cacodylate buffer (pH 7.2), postfixed for 12 h in buffered 1% osmium tetroxide, washed again in buffer, dehydrate gradea n di d serie acetonef so embedded an , Spurr'n di s epoxy resin (Spurr 1969). Ultra-thin sections were poststained with uranyl acetate/lead citrat examined ean d wit Phillipha s CM12 transmission electron microscope. Results Macroscopy After 7 days, all wounded culms showed a dark, narrow, round discoloration along the wound, both on the epidermis and more noticeably on the inner culm side. With tim discoloratioe eth n expanded epidermie axiallth n yo s variably, e.gn i . Phyllostachys as a short, thin strip and in Chusquea over the whole internode.
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