Patterns of Crystal Distribution in the Woods Of

Home , Aglaia cucullata, Chisocheton cumingianus, Cipadessa, Lansium, Melia (plant)

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).

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156 IAWA Journal, Vol. 24 (2), 2003 Negi, Gupta, Chauhan & Pal — Crystals in Meliaceae 157

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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 showing 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.

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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.

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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.

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Crystals in chambered fibrous elements — Crystals in chambered fibres were observed in a few species of Meliaceae. Cipadessa baccifera (syn. C. fruticosa) contains prismatic 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 mentioned 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 crystals.

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 sometimes 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. 11 & 12).

DISCUSSION

Although crystal distribution in Meliaceae from India has been reported by many earlier workers, many deviations were found, either with regards to position of crystals, details of chambered/non-chambered and type of ray cells with crystals. Our study revealed that in all the three species of Swietenia and two species of Xylocarpus, crystals were present in non-chambered enlarged parenchyma and ray cells (idioblasts). In Toona ciliata, druses were observed in idioblasts of both ray and parenchyma. In Toona sinensis chambering was observed up to two divisions in both procumbent and upright cells of rays. Crystals in fibres were observed in Aglaia, Azadirachta, Chukrasia, Dysoxylum gobarum and D. malabaricum, Melia, Reinwardtiodendron, and Walsura. Nair (1991) had mentioned presence of crystals in fibres of Azadirachta indica, while Kribs (1930), Pearson and Brown (1932), Chattaway (1955) and Datta and Samanta (1983) recorded crystals in all types of axial parenchyma. Crystals were found in chambered fibres in Cipadessa baccifera and not in chambered parenchyma as mentioned by Metcalfe and Chalk (1950) and Datta and Samanta (1983). Our study reveals that in all the above species crystals are present both in axial parenchyma and also in fibres and that these fibres are not long strands of parenchyma as suggested by Metcalfe and Chalk. This is born out by the following observations: 1) Intrusive tip growth, as evident from the pointed ends. 2) Distinctly bordered pits mainly in the radial walls. 3) In all the species having crystals in fibres, no diffuse parenchyma (apotracheal) is present except in Aglaia andamanica, A. perviridis and Azadirachta indica. 4) In species where crystals are present both in fibres and parenchyma, the crystal chain is almost double in length in fibres as compared to axial parenchyma strand. In all these species crystals are < 10 in axial parenchyma while the are always > 10 in fibres.

ACKNOWLEDGEMENTS

The authors are thankful to Mr. A.K. Rana, Director, Forest Research Institute for his continuous support. They are also thankful to Mr. M. Singh for assistance in section cutting, Mr. B.M. Uniyal for typing the manuscript and Mr. P. Sharma for the preparation of the photo plates.

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REFERENCES Anonymous. 1963. Indian woods – their identification, properties and uses. Vol. 2. FRI, Manager of Publications, Delhi, India. Carlquist, S. 1988. Comparative wood anatomy. Systematic, ecological, and evolutionary aspects of dicotyledon wood. Springer-Verlag, Heidelberg, Berlin. Chattaway, M.M. 1955. Crystals in woody tissues, Part I. Trop. Woods 102: 55–74. Chattaway, M.M. 1956. Crystals in woody tissues, Part II. Trop. Woods 104: 100–124. Datta, P.C. & P. Samanta. 1983. Wood anatomy of some Indo-Malayan Meliaceae. J. Indian Bot. Soc. 62: 185–203. Gregory, M. 1994. Bibliography of systematic wood anatomy of Dicotyledons. IAWA J., Suppl. 1, Leiden, Netherlands. IAWA Committee. 1989. IAWA list of microscopic features for hardwood identification. IAWA. Bull. n.s. 10: 219–332. Jain, S.S. & S.S.R. Bennet. 1997. Meliaceae. Flora of India Vol. 4: 453–526. Kribs, D.A. 1930. Comparative anatomy of the woods of the Meliaceae. Amer. J. Bot 17: 724– 738. Metcalfe, C.R. & L. Chalk. 1950. Anatomy of the Dicotyledons. Ed.1. Clarendon Press, Ox- ford. Nair, M.N.B. 1991. Wood anatomy of some members of the Meliaceae. Phytomorphology 41: 63–73. Panshin, A. J. 1933. Comparative anatomy of the woods of the Meliaceae, subfamily Swieten- ioideae. Amer. J. Bot. 20: 638–668. Pearson, R.S. & H.P. Brown. 1932. Commercial timbers of India. Vol. 1. Central Publ. Branch, Calcutta, India. Pennington, T.D. & B.T. Styles 1975. A generic monograph of the Meliaceae. Blumea 2: 419– 540. Solereder, H. 1908. Systematic anatomy of the Dicotyledons. Vol. II. Clarendon Press, Oxford.

APPENDIX — Species and samples examined

Locality abbreviations: HiP = Himachal Pradesh O = Orissa A = Andaman K = Karnataka Pu = Punjab AP = Andhra Pradesh Ke = Kerala Ra = Rajasthan AruP = Arunachal Pradesh M = Myanmar TN = Tamil Nadu As = Assam MadP = Madhya Pradesh Ut = Uttranchal Bi = Bihar Mah = Maharastra WB = West Bengal

Species name Sample numbers Locality Aglaia andamanica DDw 7827–30, 8303 A argentea DDw 6357 M cucullata DDw 414, 4554, 5135, 6653 WB edulis DDw 7535, 7954–57 As elaeagnoidea DDw 3964, 4582, 6412, 6711, 8474 AP, K maiae DDw 4535 TN perviridis DDw 7976 As spectabilis DDw 1255, 2192, 5225–27, 5325, 5890, 5970, 6061, 6462, 6780, 7134, 7180–92, 7308, 7366, 7375, 7395, 7534 A, As, M, WB Aphanamixis polystachya DDw 711, 1261, 1362, 2331, 5902, 6353, 8363, 8414, 8544 A, As, Ke, Ut

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Azadirachta indica DDw 463, 1053, 3910, 5788 AP, Ra Chisocheton cumingianus DDw 6672, 8235 AruP, M divergens DDw 6779 M nicobarianus DDw 6148, 6240 M Chukrasia tabularis DDw 1260, 1401, 2197, 2516, 5577, 5842, 6850, 7316, 7846, 7321, 7345, 7376, 7421, 7507, 7515, 7531, 7564, 7845, 7847 As, K, M, WB velutina DDw 6669, 7101 M Cipadessa baccifera DDw 3499, 3831, 6717 O Dysoxylum alliarum DDw 1259, 7244, 7377, 7536, 7571 As binectariferum DDw 5653, 6209, 6276, 6647, 8359 As, K, WB ficiforme DDw 4623 TN gobarum DDw 6447 M malabaricum DDw 4534, 5800, 5849, 5956, 6028, 6122, 6715, 6721, 6783 K, Ke, TN Khaya senegalensis DDw 8072 Mah Melia azedarach DDw 145, 946, 1201, 5803 HiP, Pu birmanica DDw 4897, 6553 M dubia DDw 705, 1093, 5660, 6552, 8329, 8373, 8456 As, K, TN, WB Reinwardtiodendron anamalaiense DDw 6720 K Sandoricum koetjape DDw 804, 5147, 6495 As, M Soymida febrifuga DDw 194, 1123, 1240, 2113, 3566, 3824, 4060, 5751, 7836, 8044 AP, K, MadP, O Sphaerosacme decandra DDw 3392 WB Swietenia humilis DDw 6707 WB macrophylla DDw 3923, 6706, 6726 Ke, WB mahagoni DDw 1361, 4568, 6708, 6725, 7471 Bi, Ke, WB Toona ciliata DDw 8, 177, 272, 712, 763, 803, 1054, 1191, 1229, 1266, As, K, MadP, 3476, 4437, 6291, 6323, 6567, 7103, 7253, 7267 Mah, Ut, WB microcarpa DDw 655, 3623, 7252, 7332, 7562, 7569 WB sinensis DDw 25, 430, 782, 897, 920, 4486, 5867 Ut Trichilia connaroides DDw 3459, 6208, 6587 Bi, M Xylocarpus granatum DDw 2239, 6262, 6609, 7557, 7591 A, WB moluccensis DDw 2514, 6269, 6306, 6819 M Walsura glauca DDw 6621 M robusta DDw 1986, 8298, 8370 A, As trifolia DDw 4232, 4627, 8358 AP, Ke, TN villosa DDw 6284, 6534, 7125, 8408 A, M

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