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Comparative Anatomy of Resin Ducts of the Pinaceae

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Comparative Anatomy of Resin Ducts of the Pinaceae ___________________________________________________________________________________www.paper.edu.cn Trees (1997) 11: 135–143 Springer-Verlag 1997 ORIGINAL ARTICLE Hong Wu c Zheng-hai Hu gomprtive ntomy of resin duts of the inee Received: 12 September 1995 / Accepted: 14 March 1996 AbstractmResin ducts are common in the Pinaceae. The these studies are limited to a specific genus or organ. A comparative anatomy of stems and leaves of 50 species and systematic investigation on the comparative anatomy of two varieties from ten genera has been investigated. The resin ducts of Pinaceae has not been undertaken. Previous structure and distribution of resin ducts differ among studies have shown that the structure and distribution of genera. Resin ducts occur in foliage leaves of ten genera resin ducts are different between different genera; therefore, of Pinaceae. Cortical resin ducts are absent in the stems of comparative studies on resin ducts of the Pinaceae have Pseudolarix and Larix. Resin ducts only occur in the great significance in theory. The present paper systemati- secondary xylem of stems of Pinus, Picea, Cathaya, cally compares the structure and distribution of resin ducts Larix, Pseudotsuga and some Keteleeria species. All of in the stems and leaves of ten genera of Pinaceae and the epithelial and sheath cells are alive and thin-walled in discusses generic relationships. the resin ducts of stem cortex and mesophyll. Except for Pinus the epithelial cells of resin ducts in the secondary xylem of stems have thick, lignified walls. Comparative study shows there are obvious differences in the resin ducts of different genera; apparent differences do not exist, wterils nd methods however, in the resin ducts of different species of the Experimental materials included 50 species and two varieties of ten same genus. According to the structure and distribution of genera of Pinaceae. Species and locations are as follows: the resin ducts in ten genera of Pinaceae, a synoptical arrangement of the genera is given and generic relation- Genus Species Locations ships of the Pinaceae are discussed. 1. Keteleeria Carr. K. cyclolepis Flous Guangxia K. davidiana (Franch.) Beiss. Zhejianga K. evelyniana Mast Yunnana c Key wordsmComparative anatomy c Pinaceae Resin duct K. fortunei (Murr.) Carr. Jiangxi 2. Abies Mill. A. chensiensis v. Tiegh Shaanxi A. ernesti Rehd. Sichuana sntrodution A. fabri (Mast.) Craib Jiangxi A. fargesii Franch Shaanxi A. faxoniana Rehd. et Wils. Sichuana The Pinaceae is the largest family of Gymnosperms with A. firma Sieb. et Zucc. Jiangxi ten genera and 230 species distributed all over the world; A. georgei Orr Yunnana there are ten genera and approximately 90 species in China A. holophylla Maxim. Jilina (Cheng et al. 1978). Some information has been reported on A. nephrolepis (Trautv.) Maxim. Shaanxi the structure, distribution and development of resin ducts in A. sutchuenensis (Franch.) Shaanxi Rehd. et Wils. the Pinaceae (Hanes 1927; Mergen and Echols 1955; Werker and Fahn 1969; Fahn 1979; Wu et al. 1987; Wu 3. Pseudotsuga Carr. P. gaussenii Flous Jiangxi 1990). A few references are also available on the compara- P. sinensis Dode Jiangxi tive anatomy of the resin ducts (Bannan 1936; Greguss 4. Tsuga Carr. Sect. Heopeuca Kang et Keng f. Guang- 1955; Gui and Lee 1963; Lee and Mu 1966; Jain 1976), but T. longibracteata Cheng donga Sect. Tsuga T. chinensis (Franch.) Pritz Shaanxi T. chinensis var. tchekiangensis Jiangxi Hong Wu ( ) c Zheng-hai Hu (Flous) Cheng et L. K. Fu Institute of Botany, Northwest University, Xi’an 710069, People’s T. dumosa (D. Don) Eichler Yunnana Republic of China T. forrestii Downie Yunnana ___________________________________________________________________________________中国科技论文在线 www.paper.edu.cn 136 Genus Species Locations distribution of the resin ducts of different genera of Pina- ceae are different. 5. Cathaya Chun et C. argyrophylla Chun et Kuang Sichuan Kuang 6. Picea Dietr. Sect. Picea Structure and distribution of the resin ducts of each P. abies (Linn.) Karst Jiangxi Pinaceae genus P. asperata Mast Shaanxi P. crassifolia Kom. Ninxiaa P. koraiensis Nakai Jilina Keteleeria Carr. P. wilsonii Mast Jiangxi Sect. Casicta Mayr The resin ducts are distributed in stem cortex and meso- P. likiangensis (Franch.) Pritz. Yunnana P. purpurea Mast. Qinhara phyll. Cortical ducts occur in the central portion of the Sect. Omerica Willk cortex (Fig. 1). In the needles, two lateral, abaxial ducts P. brachytyla (Franch.) Pritz Shaanxi appear in the mesophyll, one each on both sides of the 7. Larix Mill. L. chinensis Beissn Shaanxi vascular bundle paralleling the long axis of the leaf (Fig. 3). L. gmelini (Rupr.) Rupr. Heilong- Both epithelial and sheath cells are alive and thin-walled. In jianga addition, vertical resin ducts occur in the secondary xylem L. griffithii Hook. f. Yunnana of stems in K. davidiana (Fig. 2) and K. evelyniana. Their L. kaempferi (Lamb.) Carr. Jiangxi 2 L. mastersiana Rehd. Sichuana distribution density is 0.37 –2.5 ducts/mm . Almost all L. olgensis Henry var. koreana epithelial and sheath cells have thickened, lignified walls. Nakai Jilina L. potaninii Batal. Sichuana L. principis-rupprechtii Mayr Jiangxi L. sibirica Ledeb. Xinjianga Abies Mill. 8. Pseudolarix P. amabilis (Nelson) Rehd. Jiangxi The resin ducts are distributed in stem cortex and meso- Gord. phyll. Cortical ducts occur in the central portion of the 9. Cedrus Trew C. deodara (Roxb.) Loud. Shaanxi cortex (Fig. 4). In the needles, there are either two resin 10. Pinus Linn. Subgen. Strobus (Sweet) Rehd. ducts (A. chensiensis, A. ernestii, A. fargesii, A. faxoniana, P. armandi Franch. Shaanxi A. georgei, A. holophylla, A. nephrolepis, A. sutchuenensis) P. bungeana Zucc. ex Endl. Shaanxi or four resin ducts (A. fabri, A. firma; Fig. 5). In the former, P. koraiensis Sieb. et Zucc. Liaonina Subgen. Pinus two lateral, abaxial or middle ducts appear in mesophyll; in P. densiflora Sieb. et Zucc. Liaonina the latter, four ducts, divided into groups of two, are P. elliottii Engelmann Hainan distributed on both sides of vascular bundle. Both epithelial P. finlaysoniana Wall. Hainan and sheath cells are alive and thin-walled. P. massoniana Lamb. Sichuana P. tabulaeformis Carr. Shaanxi P. taiwanensis Hayata Anhuia P. thunbergii Parl Zhejianga Pseudotsuga Carr. P. yunnanensis Franch. Yunnana a Collected from herbarium of WNU and WUG The resin ducts are distributed in the cortex, primary xylem, and secondary xylem of stems and in the mesophyll. Cortical ducts occur in the outer portion of cortex (Fig. 6). Fresh material of 26 species was directly fixed in FAA (formalin- acetic acid-alcohol). Herbarium material of 26 other species was steeped in distilled water for several days, then fixed in FAA. Leaves, R including parts from tip, median, and base, and juvenile stems were Figs. 1–12m1 Cross-section of juvenile stem of Keteleeria fortunei dehydrated through an alcohol series and embedded in paraffin wax. showing distribution of cortical ducts. 2 A cross-section of wood of K. The samples were sectioned at 8–12 µm thickness. Wood from tree davidiana showing distribution of vertical ducts of the secondary trunks was boiled in water for 2 h, then softened in glycerine and 95% xylem. 3 A cross-section of leaf of K. fortunei showing distribution alcohol (1:1) for several weeks. Cross, radial and tangential sections of of ducts in mesophyll (arrow). 4 A cross-section of juvenile stem of the wood were obtained with a sliding microtome at 20–40 µm Abies fabri showing distribution of cortical ducts. 5 A cross-section of thickness. Sections were stained with safranin and fast green, sealed leaf of A. fabri showing distribution of ducts in mesophyll (arrows). in Canada balsam, and observed under an Olympus microscope. 6 A cross-section of juvenile stem of Pseudotsuga sinensis, showing distribution of cortical ducts. 7 Cross-section of wood of P. sinensis showing distribution of vertical ducts of the secondary xylem. 8 A longitunal (tangential) section of wood of P. sinensis showing distri- bution of horizontal ducts of the secondary xylem. 9 A cross-section of esults leaf of P. sinensis showing distribution of ducts in mesophyll (arrow). 10 A cross-section of juvenile stem of Tsuga chinensis showing The resin ducts are schizogenously formed in the stems and distribution of cortical ducts. 11 A cross-section of leaf of T. chinensis leaves of the Pinaceae. Mature ducts are elongated struc- showing distribution of duct in mesophyll (arrow). 12 A cross-section of juvenile stem of Cathaya argyrophylla showing distribution of tures consisting throughout their length of epithelial cells cortical ducts. Bars indicate 500 µm. D, Dead sheath cell; E, Epithelial surrounding an intercellular space and on their outer side of cell; HR, Horizontal duct; L, Epithelium with lignified wall; Rd, Resin one to three layers of sheath cells. The structure and duct; T, Epithelium with thin wall; VR, Vertical duct ___________________________________________________________________________________中国科技论文在线 www.paper.edu.cn 137 ___________________________________________________________________________________中国科技论文在线 www.paper.edu.cn 138 Figs. 13–23m13 A cross-section of wood of C. argyrophylla showing section of leaf of P. wilsonii showing distribution of ducts in mesophyll distribution of vertical ducts of the secondary xylem. 14 A longitudinal (arrows). 20 A cross-section of juvenile stem of Larix kaempferi (tangential) section of wood of C. argyrophylla showing distribution of showing cyst-like vesicle (arrow) in cortex near phloem. 21 A cross- horizontal ducts of the secondary xylem. 15 A cross-section of leaf of section of wood of L. kaempferi showing distribution of vertical ducts C. argyrophylla showing distribution of ducts in mesophyll (arrows). of the secondary xylem. 22 A longitudinal (tangential) section of wood 16 A cross-section of juvenile stem of Picea wilsonii showing of L. kaempferi showing distribution of horizontal ducts of the distribution of cortical ducts. 17 A cross-section of wood of P. wilsonii secondary xylem. 23 A cross-section of leaf of L.
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