Patterns of Crystal Distribution in the Woods Of
Total Page:16
File Type:pdf, Size:1020Kb
IAWA Journal, Vol. 24 (2), 2003: 155–162 PATTERNS OF CRYSTAL DISTRIBUTION IN THE WOODS OF MELIACEAE FROM INDIA by Krishna Negi, Sangeeta Gupta, Luxmi Chauhan & Mohinder Pal Wood Anatomy Discipline, Forest Research Institute, P.O. New Forest, Dehra Dun, India SUMMARY A detailed wood anatomical survey of 42 species of Meliaceae from India has been made with a view to study the distribution pattern of different types and location of crystals. Interestingly, all the IAWA features of prismatic crystals were found in this family. The confusion regarding presence of crystals in fibres or axial parenchyma strands is clarified. Key words: Crystals, druses, fibres, Meliaceae. INTRODUCTION Information on the occurrence of crystals in plant cells of the dicotyledons has been summarised by Solereder (1908), Kribs (1930), Metcalfe and Chalk (1950), Chattaway (1955, 1956) and Carlquist (1988). Panshin (1933) and Pennington and Styles (1975) have studied Meliaceae in particular. For woods of Meliaceae from India, Pearson and Brown (1932), Anonymous (1963), Datta and Samanta (1983) and Nair (1991) have published detailed anatomical descriptions of many species of this family. However, a comprehensive account of the distribution of crystals in Meliaceae from India is lacking. Crystals are diagnostic in wood identification, owing to their distinctive appearance, location, and type (Chattaway 1955). During the course of wood anatomical studies of Meliaceae from India, the presence of crystals in 42 species of 18 genera was noted. MATERIAL AND METHODS The basis of this study was formed by 216 authentic wood samples of 42 species in 18 genera, housed in the Xylarium of the Wood Anatomy Discipline, Forest Research Institute, Dehradun (see Appendix). The nomenclature given by the Flora of India (Jain & Bennet 1997) has been followed. Cross, tangential and radial sections were examined for the occurrence of crystals. Small blocks of the wood were boiled for softening and sections of 20 µm were cut. Unstained sections were bleached and permanently mount- ed for recording the presence or absence of crystals. Small radial chips were macerated following Schultzʼs method, i.e. 30% nitric acid and a pinch of potassium chlorate. The terminology followed in describing the crystal types is as given by Chattaway (1955, 1956), Carlquist (1988) and IAWA (1989). Downloaded from Brill.com10/02/2021 01:17:48PM via free access 156 IAWA Journal, Vol. 24 (2), 2003 Negi, Gupta, Chauhan & Pal — Crystals in Meliaceae 157 ➙ ➙ 1 2 3 4 5 Fig. 1. Azadirachta indica: transverse section (TS) showing crystals in parenchyma and fibres. – Fig. 2. Dysoxylum gobarum: radial longitudinal section (RLS) showing chambered crystals in fibre (left) and parenchyma (right). – Fig. 3. Azadirachta indica: macerated material show- ing chambered crystals in fibre. – Fig. 4. Swietenia mahagoni: RLS showing small crystals in upright ray cells. – Fig. 5. Cipadessa baccifera: tangential longitudinal section (TLS) showing lacuna in the centre of crystals. — Magnification of all× 400. Downloaded from Brill.com10/02/2021 01:17:48PM via free access 156 IAWA Journal, Vol. 24 (2), 2003 Negi, Gupta, Chauhan & Pal — Crystals in Meliaceae 157 Table 1. Patterns of crystal distribution in Indian Meliaceae. Species Fibres URC PRC Parenchyma Crystals No. of No. of C NC C NC C NC variable CCF CCP Aglaia andamanica + + 15–26 4–6 argentea + + 15–25 4–6 cucullata + + 15–20 4–5 edulis + + 15–20 4–8 elaeagnoidea + + 15–25 4–5 maiae + + 15–25 4–6 perviridis + + 15–20 4–6 Azadirachta indica + + 15–20 4–8 Chisocheton cumingianus + 4–6 divergens + 4–8 nicobarianus + 4–5 Chukrasia tabularis + + 17–20 4–8 velutina + + 15–18 4–7 Cipadessa baccifera + 12–20 Dysoxylum alliarium + 4–6 binectariferum + 4–8 ficiforme + 4–6 gobarum + + 15–20 4–6 malabaricum + + 15–20 4–8 Khaya senegalensis +(i) Melia azedarach + + 13–14 4–8 birmanica + + 12–15 4–8 Reinwardtiodendron anamalaiense + + 18–20 4–6 Soymida febrifuga + +(i) +(i) Swietenia humilis +(i) + + + macrophylla +(i) + + + mahagoni +(i) + + + Toona ciliata +(i) +(i) 2–5 sinensis +(i) +(i) Trichilia connaroides + 4–5 Xylocarpa granatum +(i) + +(i) moluccensis +(i) + +(i) Walsura glauca + + 15–20 2–6 robusta + + 15–20 4–7 trifolia + + 16–29 4–8 villosa + + 15–20 4–6 URC: upright ray cells; PRC: procumbent ray cells; C = chambered, NC = non-chambered. + = crystals present; (i) = idioblasts present. CCF: crystals in chambered fibres; CCP: crystals in chambered parenchyma. OBSERVATIONS Table 1 summarises the crystal distribution. All the IAWA features of prismatic crystals were found in this family. Crystals were absent in 6 species, viz. Aglaia spectabilis, Aphanamixis polystachya, Melia dubia, Sandoricum koetjape, Sphaerosacme decandra, and Toona microcarpa. The following patterns of crystal distribution were observed in the remaining species. Downloaded from Brill.com10/02/2021 01:17:48PM via free access 158 IAWA Journal, Vol. 24 (2), 2003 Negi, Gupta, Chauhan & Pal — Crystals in Meliaceae 159 ➙ 7 ➙ 6 8 9 10 11 12 Fig. 6. Xlyocarpus granatum: RLS showing crystals in non-chambered upright and procumbent ray cells. – Fig. 7. Toona sinensis: RLS showing crystals in chambered upright ray cells. – Fig. 8. Toona sinensis: RLS showing crystals in chambered procumbent ray cells. – Fig. 9. Swietenia macrophylla: RLS showing crystals in non-chambered axial parenchyma. – Fig. 10. Chisocheton cumingianus: RLS showing crystals in chambered parenchyma cells. – Fig. 11. Toona ciliata: RLS showing druses in enlarged (idioblast) ray cell. – Fig. 12. Toona ciliata: RLS showing druses in chambered parenchyma cells. — Magnification of all× 400. Downloaded from Brill.com10/02/2021 01:17:48PM via free access 158 IAWA Journal, Vol. 24 (2), 2003 Negi, Gupta, Chauhan & Pal — Crystals in Meliaceae 159 Crystals in chambered fibrous elements — Crystals in chambered fibres were observed in a few species of Meliaceae. Cipadessa baccifera (syn. C. fruticosa) contains pris- matic crystals in chambered fibres while they are absent in axial parenchyma and rays. Some crystals of this species have lacuna in the centre of the crystals (Fig. 5) as men- tioned by Carlquist (1988). Moreover, the size of the crystals is uniform throughout the length of the fibre. Crystals in chambered fibres were also commonly found in Aglaia andamanica, A. edulis, Azadirachta indica (Fig. 1 & 3), Dysoxylum malabaricum, D. gobaram (syn. D. procerum) (Fig. 2), Melia azedarach, M. birmanica, and Walsura trifolia (syn. W. pisicida), while they are uncommon to rare in Aglaia cucullata, A. perviridis, Chukrassia tabularis, C. velutina, Reinwardtiodendron anamalaiense (syn. Lansium anamallayanum), Walsura robusta and W. villosa. In Aglaia argentea few crystals bear a line mark in the centre while in Walsura robusta there are lacuna in the centre of the few crystals. These crystalliferous fibres frequently contain a mixture of large and small crystals. Crystals in non-chambered ray cells — In Khaya senegalensis, Soymida febrifuga, Swietenia humilis, S. macrophylla, S. mahagoni, Xylocarpus granatum (syn. Carapa obovata) and X. moluccensis (syn. Carapa moluccensis) crystals are present in both procumbent ray cells and upright ray cells (Fig. 6). Upright ray cells that have crystals are idioblasts while the procumbent cells containing crystals are normal in size. In the three species of Swietenia, prismatic crystals characteristically appear in two forms, solitary and 2–3 crystals in non-chambered cells. These crystals are smaller in size and seem to be crystal fragments of different shapes (Fig. 4). Sometimes crystals have a line mark on their surface and sometimes a lacuna is present in the centre of the crys- tals. Crystals in chambered ray cells — The upright and procumbent ray cells of Toona sinensis (syn. T. serrata) possess prismatic crystals. Sometimes these crystal-bearing cells are idioblasts and sometimes they are subdivided into not more than two chambers (Fig. 7 & 8). Crystals in non-chambered parenchyma — In Soymida febrifuga, Swietenia humilis, S. macrophylla, S. mahagoni, Xylocarpus granatum and X. moluccensis crystals are mostly present in non-chambered parenchyma cells (Fig. 9). In Xylocarpus granatum and X. moluccensis prismatic crystals are present in slightly enlarged, non-chambered parenchyma cells. In Soymida febrifuga, crystals are present in idioblasts. Crystals in chambered parenchyma — Crystals in chambered parenchyma are present quite frequently in Aglaia andamanica, A. argentea, A. edulis, Azadirachta indica, Chisocheton cumingianus (syn. C. paniculatus), C. divergens, C. nicobrianus (syn. C. grandiflorus), Chukrasia tabularis, C. velutina, Trichilia connaroides (syn. Heynia trijuga), Dysoxylum gobarum (Fig. 2), D. malabaricum, Melia azedarach, and Wal- sura trifolia, while they are very infrequent in Aglaia cucullata, A. maiae, A. elaeag- Downloaded from Brill.com10/02/2021 01:17:48PM via free access 160 IAWA Journal, Vol. 24 (2), 2003 Negi, Gupta, Chauhan & Pal — Crystals in Meliaceae 161 noidea, A. perviridis, Dysoxylum alliarum (syn. D. hamiltonii), D. binecteriferum, D. ficiforme (syn. D. purpureum), Melia birmanica, Reinwardtiodendron anamalaiense, Walsura glauca, W. robusta, and W. villosa (Fig. 10). In Dysoxylum malabaricum the axial parenchyma strands which bear crystals some- times show further vertical division thereby making two small chambers containing crystals. Druses — Only Toona ciliata has druses in idioblasts in chambered and non-chambered parenchyma cells and also in upright ray cells (Fig.