Development of Woody Branch Attachments in Schefflera (Araliaceae Or Apiaceae) Author(S): P
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Development of Woody Branch Attachments in Schefflera (Araliaceae or Apiaceae) Author(s): P. Barry Tomlinson, Jack B. Fisher, Francis Hallé, Randol Villalobos Reviewed work(s): Source: American Journal of Botany, Vol. 92, No. 11 (Nov., 2005), pp. 1765-1773 Published by: Botanical Society of America Stable URL: http://www.jstor.org/stable/4125506 . Accessed: 04/04/2012 11:58 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Botanical Society of America is collaborating with JSTOR to digitize, preserve and extend access to American Journal of Botany. http://www.jstor.org American Journalof Botany 92(11): 1765-1773. 2005. DEVELOPMENT OF WOODY BRANCH ATTACHMENTS IN SCHEFFLERA (ARALIACEAE OR APIACEAE)' P. BARRY TOMLINSON,2,6 JACK B. FISHER,3 FRANCIS HALLE*,4AND RANDOL VILLALOBOS5 2HarvardForest, HarvardUniversity, Petersham,Massachusetts 01366 USA and The Kampong of the National Tropical Botanical Gardens, 4013 Douglas Road, Coconut Grove, Florida 33133 USA; 3FairchildTropical Botanic Garden, 11935 Old Cutler Road, Coral Gables, Florida 33156 USA and Departmentof Biological Sciences, Florida InternationalUniversity, Miami, Florida 33199 USA; 4109 Ave. de Lodbve, Montpellier,34070 France; and 5Departmentof Biology, University of Miami, Coral Gables, Florida 33144 USA Attachmentof branches in Schefflera is unusual in that it involves fingerlike woody extensions that originate in the cortex and pass gradually into the woody cylinder of the parent shoot. We tested the hypothesis that these structurescould be roots since Schefflera is a hemi-epiphyte with aerial roots. These branch traces originate by secondary development in the many leaf traces (LTs) of the multilacunarnode together with associated accessory traces. In the primary condition, the LTs may be described as cortical bundles. Leaves are long persistent and can maintaina primarystem connection across a broad cylinder of secondary xylem. Under the stimulus of branch development, the LTs form a template for secondary vascular development. Because the LT system is broad, with many traces, the branch attachmentis also broad. The fingerlike extensions are attached to the surface of the woody cylinder of the parent stem but are progressively obscured as a continuous cambium is formed. Bark tissues are included within the branch axil because of the extended cortical origin of the initial attachment.The results are discussed in the context of branch-trunkunions in tropical plants, an importantcomponent of canopy development. Key words: Araliaceae; branch insertion; leaf gap; leaf trace; nodal anatomy;Schefflera; secondary xylem. There is little information concerning the structuralmech- axis. He hypothesized that these branches (or "reiterations") anisms involved in secondary changes while woody branches might represent roots extending from the base of the branch develop a vascular connection to their parent axis. This omis- similar to the way aerial roots develop in this and some other sion is largely a consequence of the need to study large struc- woody species (Hall6, 1991). However, no developmentalev- tures at a microscopic level. Standardtexts may devote only idence was offered. We show that quite complex processes are a few paragraphsto the topic (e.g., Fink, 1999; Larson, 1994). involved in the productionof these fingerlikeextensions of the In contrast, there are extensive descriptions of nodal vascula- branch. The primary configurationis itself complex, and the ture in dicotyledons, in which primary constructionmay have result may be interpretedas a particularlyefficient method of systematic relevance (Howard, 1979), but they relate largely branchinsertion. To the best of our knowledge, this distinctive to leaf traces (LTs) alone, with incidental reference, if at all, developmentalprocess has not been reportedpreviously. Using to branch traces in the primary condition (Eames and Mac- developmental evidence, we tested an initial hypothesis that Daniels, 1947; Esau, 1965, 1977). The chief emphasis is on leaf traces and branch traces are associated in unusual ways. herbaceous plants and simple systems where bud traces are attachedto the margins of stelar leaf gaps, but without details MATERIALSAND METHODS of how an axillary bud develops a mature stem attachment. Branch-trunkattachments are known to be fea- Two species of Schefflera were studied; one tree species, S. actinophylla important (Endl.) Harms, is and can be invasive because of its a tures of structuralcompartmentalization in tree construction widely planted habit; smaller species, S. arboricola (Hayata) Merr., is a commonly cultivated or- (Shigo, 1985) and are certainly an essential component in the of a forest because branch namental shrub. We examined S. arboricola in greater detail because of its development canopy persistence smaller size, with the axes of the order of 1 cm which can demonstratesmechanical and primary diameter, physiological efficiency. Knowl- be conveniently serially sectioned. edge of branchattachment is also importantin correctpruning practices in arboriculture(Shigo, 1989). We draw attentionto Surface morphology--Fresh or partly dried stems can be decorticatedby this topic by describing the distinctive method of branch at- removing all tissues external to the cambium to reveal the branchattachment tachment in the tropical genus Schefflera (Araliaceae or Api- with reference to the surface morphology of the secondary xylem of the parent aceae, Judd et al., 2002). The leaves are sheathing, with many axis (Figs. 1-7). Some of these specimens were subsequently sawn trans- LTs arising from a multilacunarnode (Howard, 1979). versely (Fig. 4). Schefflera can develop as a hemi-epiphyte producing aerial anatomical material was either unfixed roots. Hall6 (1991) illustratedunusual fingerlike extensions of Anatomy-For study, or fixed in wood from lateral branches where they connect with the main formalin-acetic-alcohol(FAA; 85 parts 70% ethanol, 10 parts glacial acetic acid, and 5 parts 40% formaldehyde) and sectioned on a sliding microtome. Fixed materialwas washed well in tap water before Sections were ' Manuscriptreceived 25 March 2005; revision accepted 2 August 2005. sectioning. cut and to a lesser extent at 60-120 This researchwas supportedby the Eleanor Crum Professorshipin Tropical transversely longitudinally [m thickness, or to ensure Botany of the National TropicalBotanical Garden. Facilities at the Kampong serially sequentially, whole sections for vascularcontinuity. Sec- of the National Tropical Botanical Garden provided by its Director, Larry tions were all stained in 0.5% aqueous toluidine blue (Sigma, St. Louis, Mis- Schokman, are greatly appreciated. souri, USA) and mounted in glycerine : water (1 : 1) as temporaryprepara- 6 Author for correspondence(e-mail: [email protected]) tions. To facilitate photographic documentation, selected sections were 1765 1766 AMERICAN JOURNAL OF BOTANY [Vol. 92 Figs. 1-7. Schefflera spp., decorticateddried axes. Figs. 1, 2. S. cf. actinophylla. Figs. 3, 4. S. actinophylla. Figs. 5-7. S. arboricola. 1. Branch insertion from the side; orientationsare changed; the branch was originally vertical and the main axis oblique. 2. Front view of Fig. 1. 3. Older axis showing continuous complete fusion of branchbase with parent axis, arrows indicate acropetalconnections. 4. Transversesection of an older axis showing inclusions of cortex and secondary phloem between enveloping branch and parent axis with collapsed pith. 5. Axis with branches attached at three successive nodes below an aborted trunk. Arrows indicate long median branch traces. 6. Older axis from front with more complete fusion. Arrows indicate scars of leaf traces. 7. Branch insertion below remains of inflorescence (infl), branch traces almost completely fused; arrow as in Fig. 5. Scale bars = 4 cm for Figs. 1-4; = 1 cm for Figs. 5-7. November 2005] TOMLINSONET AL.-WOODY BRANCHESIN SCHEFFLERA 1767 bleached for 10-20 min in sodium hypochlorite (commercial bleach) diluted cular bundles separatedby narrowrays (Figs. 8-11). Each vas- 1 : 2 with tap water to remove cell contents. Such sections were rinsed well cular bundle is delimited externally by a strandof thick walled in tap water, dehydratedin ethanol, and stained in 1% safranin (Sigma) and fibers seen in transverse section (TS) as a cap (see f in Figs. 1% alcian green (Sigma Chemical, St. Louis, MO) (5 : 1 mix) both in 70% 11, 12). Internally,LTs close to their departurefrom the stele ethanol and mounted in Permount (Fisher Scientific, Fair Lawn, New Jersey, are recognized by extensively developed primaryxylem (Figs. USA). 9, 10). Such exiting LTs leave a narrow gap in the parent To the axes with little were study primary vasculature, secondary growth vascular cylinder (lg in Fig. 11), the capable of subsequent followed axes with branch attachmentsof gap initially sectioned, by developing complex development as describedlater (in Basal attachment). size, to when the branch attachmenthad become increasing continuing stages Cortical features include a well-developed collenchymatous quite woody. Because