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1980 43 CYSTOLITHS in the SECONDARY XYLEM of SPARATTANTHELIUM (HERNANDIACEAE) by Ben JH Te

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1980 43 CYSTOLITHS in the SECONDARY XYLEM of SPARATTANTHELIUM (HERNANDIACEAE) by Ben JH Te lAW A Bulletin n.s., Vol. 1 (1-2), 1980 43 CYSTOLITHS IN THE SECONDARY XYLEM OF SPARATTANTHELIUM (HERNANDIACEAE) by Ben J. H. ter Welle Institute of Systematic Botany, University of Utrecht, Heidelberglaan 2, Utrecht, The Netherlands Abstract Cystoliths were observed in the secondary the leaves of Ficus. As cystoliths have also been xylem of Sparattanthelium (Hernandiaceae). described for the leaves of Sparattanthelium Their shape, size, distribution and chemical (Solereder, 1899) and have been accepted as a composition is described. The systematic value feature of systematic value (e.g. Kubitzki, of cystoliths in the Hernandiaceae as well as in 1969), a supplementary examination of the general is discussed. wood of other species was undertaken. Introduction Methods Cystoliths are internal, stalked outgrowths Both stained and unstained sections were of the cell wall that project into the cell lumen. studied. A double staining of Astrablue and They consist of cellulose and are impregnated Safranin was applied to differentiate between with calcium carbonate, they are irregular in unlignified and lignified cell components. The shape and sometimes fill up the cell completely. usual light-microscope, polarized-light and SEM­ Cystoliths may occur in parenchymatous cells techniques were used. Finally, the mineral in various parts of the plant, including even composition of the cystoliths was determined xylem and phloem rays. They are most fre­ with an Energy Dispersive X-ray Analyzer. quently found in the epidermis, in hairs or in special large cells which are termed litho cysts Materials (Fahn, 1967). In spite of the presence of cal­ According to Kubitzki (1969), the genus cium carbonate, this type of cell inclusions was Sparattanthelium comprises 13 species, lianas not included in the comprehensive study on or shrubs, restricted to the neotropics. For this the occurrence and shape of crystals made by study wood samples of 5 species were available. Chattaway (1955, 1956). Others, like Metcalfe In addition samples of other Hernandiaceae & Chalk (1950), Esau (1963) and Fahn (1967) and of a few taxa of the Opiliaceae and of the also described cystoliths as a separate category, Acanthaceae were investigated. apart from crystals. The occurrence of cystoliths in various parts Material seen: of the plant, but especially in the leaves, is re­ Hernandiaceae - Gyrocarpus americanus stricted to a few families (Metcalfe & Chalk, Jacq. ssp. americanus, FHOw 70 I, Sri Lanka; 1950; Pireyre, 1961), particularly in Cannabi­ Pulle 3217, New Guinea; Hernandia didymantha naceae, Moraceae and Urticaceae. Donn., Uw 10383, P.H. Allen, Costa Rica, ex Cystoliths in the secondary xylem were up USw 30169; H. guianensis Aubl., Uw 380, till now only known in a single family, Opilia­ Stahel 380, Suriname; Uw 1509, Lanjouw & ceae (Record, 1925, 1927; Kanehira, 1921; Lindeman 1547, Suriname; H. peltata Meissn., Metcalfe & Chalk, 1950). Record described B.W. 7670, New Guinea; Illigera pentaphylla cystoliths as 'deposits of calcium carbonate in Welw., Uw 22123, Versteegh & den Outer 588, special structures in the rays of 7 genera, viz. Ivory Coast; Sparattanthelium aruakorum Agonandra, Cansjera, Champereia, Lepionurus, Tutin, Uw 12148, van Donselaar 3754, Suri­ Meliantha, Opilia and Rhopalopilia'. According name; S. borororum Mart., Uw 24192, R.C. to Metcalfe & Chalk (1950), Agonandra is the Gill 43, Equador, ex SJRw 36125; S. glabrum only genus which lacks the cystoliths. Rusby, Uw 24073, Schunke 2539, Peru, ex By chance I observed in a section of Sparat­ USw 41876; S. guianense Sandw. Uw 24087, tanthelium deposits which looked like a bunch Fanshawe, F.D. 3938, Guyana, ex FHOw of grapes. At first sight, these deposits appear­ 13266; S. wonotoboense Kosterm., Uw 24191, ed to be similar to the well-known cystoliths in A.C. Smith 3390, Guyana, ex SJRw 35917. Downloaded from Brill.com09/26/2021 07:44:50AM via free access 44 IAWA Bulletin n.s., Vo1.1 (1-2),1980 Fig. 1-9. Light-microscopical appearance of cystoliths in the secondary xylem. -- I, 2 & 3: Spa­ rattanthelium aruakorum Tutin. (Uw 12148). Cross section, x 35, x 90 & x 220 resp. -- 4: Sparat­ tan thelium guianense Sandw. (Uw 24087). Tangential section, x 35. -- 5: Sparattanthelium wono­ toboense Kostenn. (Uw 24191). Radial section, x 35. -- 6 & 8: Sparattanthelium aruakorum Tutin. (Uw 12148). Tangential and radial section resp., x 220. --7: Trichantheragigantea (Humb. & Bonp1.) Steud. (Uw 5476). Radial section, x 220. -- 9: Champereia manillana Merrill. (Jacobs 7611). Radial section, x 220. Downloaded from Brill.com09/26/2021 07:44:50AM via free access IAWA Bulletin n.s., Vol. I (1-2), 1980 45 Opiliaceae - Agonandra silvatica Ducke, Uw Chemical composition - A thorough exami­ 3402, Lindeman 4995, Suriname; Uw 6806, nation of the chemical composition of the cys­ Schulz 8329, Suriname; Champereia manillana toliths is beyond the scope of this study. How­ Merrill, Jacobs 7611, Philippines; Opilia cf ever, in order to establish the cystolith-nature amentacea Roxb., Jacobs 9600, Papua New of the observed inclusions, some information Guinea; O. celtidifolia (Guill. & Perr.) End!. ex on this subject is necessary. Therefore, sections Walp., Breteler 2261, Cameroun. were double-stained with Safranin and Astrablue Acanthaceae - Aphelandra tetragona (Vah!) according to the method recommended by Von C.G. Nees ab Esenb., Uw 10932, Florschiitz & Aufsess (1973) to differentiate between lignin Maas 2431, Suriname; Trichanthera gigantea and cellulose. In both transverse and longitudi­ (Humb. & Bonpl.) Steud., Uw 200 I, Lanjouw nal sections the cystoliths are stained bright & Lindeman 3433, Suriname; Uw 5476, Geys­ blue. Nevertheless the intensity of the blue col­ kes s.n., Suriname. our shows various grades. There is no trace of red stained material in the cystoliths which in­ Results dicates the absence of lignin. If pedicels do oc­ cur, they remain uncoloured. Sparattanthelium Subsequently SEMjEDXA techniques were Cystoliths were found in all species of Spa­ used to determine the mineral components of rattanthelium examined. Their shape, size and the cystoliths. The only element identified is distribution was variable, both within a sample calcium (Fig. 14, 15 & 16). In polarized light and between different samples. the cystoliths show little or no birefringence. Shape - Slightly oval and oblong in S. aruakorum, S. glabrum and S. guianense. In The SEM investigations gave some indication for the presence of a thin layer of unidentified S. borororum and S. wonotoboense only ob­ long cystoliths occurred. nature between the wall of the lithocyst and the cystoliths (Fig. 13). This layer stained blue Size - The oval cystoliths measure from c. with the Astrablue-Safranin staining. Although 45 /lm up to 60 /lm and the oblong ones from cystoliths usually are not classified as crystals, 100 x 40 to 160 x 60 /lm. In S. wonotoboense, this layer is comparable with integuments de­ a cystolith of 240 x 64/lm was observed. They scribed for crystals in the secondary xylem of usually fill up the cell completely. e.g. Flacourtiaceae (Miller, 1975). Structure - The surface is always more or The occurrence of the cystoliths in Sparat­ less granular, but without the sharp points tanthelium is restricted to a certain part of the characteristic for crystals like druses. Only oc­ stem. Shutts (1960) already noticed in this casionally the cystoliths show faint radiating genus the presence of two distinct types of ves­ lines (Fig. 10). On cross sections concentric sels in one sample, indicated by diameter and lines around the centre of the cystoliths can of­ element length. On a transverse section these ten be noticed (Fig. 2 & 3). two different types are very clear. It is remark­ Distribution - Average values for the num­ able that the cystoliths often occur at the place ber of cystoliths per square mm (cross section), where the diameter of the vessels changes ab­ based on 10 measurements, were computed. ruptly. The averages vary from 2 to 29, but in general According to Kubitzki (1969), cystoliths oc­ there are less than 10 per square mm (Fig. I). cur in the leaves of all species of Sparattanthe­ Lithocysts - The shape of these cells com­ lium and Gyrocarpus, together constituting the monly resembles that of the cystoliths, since the latter fill up the cell completely. The out­ tribe Gyrocarpoideae. However, contrary to line of the lithocysts in longitudinal sections is the results obtained with the secondary xylem rounded (oval or oblong, Fig. 4 & 5). and rath­ of Sparattanthelium, no cystoliths were observ­ er irregular in cross sections (Fig. 3). ed in the wood of Gyrocarpus. In view of the fact that occurrence of cystoliths is restricted Cystolith-containing cells in Sparattanthelium to a small number of families (Solereder, 1899, are always part of a parenchyma strand, but they are much larger than the other parenchy­ 1908; Pireyrc, 1961), it seemed worthwhile to examine the wood of some families from the ma cells. In general the cells show about the leaves of which cystoliths have been reported. same size and shape as oil or mucilage cells, well known from the wood of e.g. Lauraceae. Acanthaceae Pedicel - The presence of a pedicel is of es­ Cystoliths are abundant in the ray cells of sential importance for the development of cys­ two samples examined of Trichanthera gigantea. toliths (Fahn, 1967). However, in the material The ray cells are not enlarged and similar to the examined pedicels were rarely observed and other ray cells. Cystoliths occur in both square short. and upright cells. As a result their shape varies Downloaded from Brill.com09/26/2021 07:44:50AM via free access 46 lAW A Bulletin n.s., Vol. 1 (1-2), 1980 Downloaded from Brill.com09/26/2021 07:44:50AM via free access IAWA Bulletin n.s., Vol. I (1-2),1980 47 from more or less square or round to very of­ fIrst stage of its formation, the body is incrust­ ten oblong, respectively c. 40 !.1m and from 120 ed or impregnated with amorphous calcium x 30 to 180 x 30 !.1m (Fig.
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