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Isolated Sieve Tube (S)/Elements in the Barks of Some Angiosperms IAWA Bulletin n.s., Vol. 4 (4),1983 255 ISOLATED SIEVE TUBE(S)/ELEMENTS IN THE BARKS OF SOME ANGIOSPERMS by R.R. Chavan*, J.J. Shah and K.R. Patei" Department of Biosciences, Sardar Patel University, Vallabh Vidyanagar, 388 120, Gujarat, India Summary The occurrence of solitary or groups of iso­ tures, information on naturally occurring iso­ lated tube element(s) in the interfascicular re­ lated sieve tube elements is meagre and data on gion of Aristolochia indica L., Datura innoxia their development, histochemistry and dimen­ Mil!., Lecanthus wightii Wedd. and Xanthium sional details are sparse in comparison with strumarium L. and in the secondary phloem other aspects of phloem research. The present rays of Strychnos nux-vomica L., Boehmeria study augments information in this direction. nivea Gaudieh., Leucosceptrum cannum Sm., Dahlia imperialis Roezl., Gynura angulosa DC., Materials and Methods Vernonia divergens (Roxb.) Edgew. and Verno­ The materials for the investigation were col­ nia volkameriaefolia De., encountered during lected from Central, Northeastern, Southern an investigation of the barks of 125 dicotyle­ and Western parts of India (Chavan, 1981). dons belonging to 43 families, has been record­ Five to ten sam pies of bark from the main ed. The salient developmental features of iso­ trunk at 1.5 m height of arborescent members lated sieve tube elements in the last three taxa and of internodal regions at mid-height of her­ are described involving histochemical tests and baceous members were collected from various their dimensional details. A comparative analy­ localities to randomise them and avoid specific, sis of secondary phloem of members of Com­ c1imatic and edaphic effects. They were fixed positae with and without isolated sieve tube on the spot in formalin acetic alcohol (Sass, elements indicates that taxa with a high fre­ 1958) and processed for mierotomy by conven­ quency of isolated sieve tube elements show a tional methods (Berlyn & Miksche, 1976). high incidence of compound sieve plates on Transverse, radial and tangential longitudinal predominantly very oblique end walls and also sections of 8-10 J..I.ITl thiekness were stained possess a system of broad rays. The possible with tannic acid-ferric chloride-resorcin blue signifieance of isolated sieve tube elements is combination (Cheadle et al. , 1953), safranine' discussed. '0' (Sass, 1958) and toluidine blue '0' (O'Brien Key words: Histochemistry, phloem rays, inter­ et al., 1964). Unembedded bark sampies were fascicular region, Compositae, Solanaceae, cut on a sliding microtome. Histochemical tests Loganiaceae, Urtieaceae. for total proteins, total nucleic acids, DNA, starch (Jensen, 1962) and histones (Black & Introduction Ansley, 1964) were performed. Dimensional Fischer's report on the occurrence and dis­ details were obtained with a Carl Zeiss projec­ tribution of isolated sieve tube elements and tion disc microscope and Kelner's eye piece. isolated phloem bundles in members ofCucur­ Photomicrographs were taken on a Carl Zeiss bitaceae (1884, cited in Esau, 1969) was fol­ photomicroscope with planapochromatic ob­ lowed by only a few, as in Vicia. Conium and jectives using ORWO NP-15, Kodak mono­ Aristolochia (von Guttenberg, 1943), in the chrome SO 410,35 mm black and white films. mesocarpie and endocarpie regions of Citrullus lanatus. Coccinia grandis. Luffa cylindrica and Results L. acutangula (Thanki, 1978; Shah et al., 1983) Isolated sieve tube element(s) (ISTEs) occur and in internodal regions of seven cucurbita­ either solitary or in groups, in the interfascicu­ ceous species (Karnik, 1981). Though Metcalfe lar region of Aristolochia indica. Datura inno­ and Chalk in their monumental work (1950, xia. Lecanthus wightii and Xanthium struma­ 1979) covered major aspects of anatomie al fea- rium (Figs. I & 2). In Strychnos nux-vomica. * Present address: Department of Botany, D.V.S. College of Arts and Science, Shimoga 577 201, Karnataka, India. *·Present address: School of Forestry & Environmental studies, Yale University, New Haven, Con­ necticut 06511, U.S.A. Downloaded from Brill.com09/29/2021 04:06:16AM via free access 256 IAWA Bulletin n.s., Vol. 4 (4),1983 Dahlia imperialis, Gynura angulosa, Vernonia tiation of ISTEs occurs at random in the ray volkameriaefolia, Boehmeria nivea and Leuco­ (Figs. 21, 22) frequently towards its margin sceptrum cannum solitary or isolated groups of (Figs. 8, 10). The initial of an ISTE is histologi­ sieve tube elements occur in secondary phloem cally and histochemically different from the rays (Figs. 3, 4-12, 18, 19). neighbouring cells, showing dense cytoplasm Among the Compositae Dahlia imperialis, and a prominent nucleus with higher amounts Gynura angulosa and Vernonia volkameriaefo­ of total protein, total nucleic acids, DNA, and lia, ISTEs are frequent in the wide rays show­ arginine rich histones as evidenced by mercuric ing tangentially horizontal or diagonal disposi­ bromophenol blue, azur B, Feulgen and ammo­ tions (Figs. 3-10, 14, 15, 18, 19). They may niacal silver nitrate reactions. The initial divides be solitary and isolated from the main strand parallel to its longest axis in D. imperialis and (Figs. 2, 3) or in groups of two or more, either V. volkameriaefolia (Figs. 23-28), and at right interconnected end to end to bridge adjacent angles in G. angulosa (Fig. 21) and further the main secondary phloem strands (Figs. 5, 10, 11, derivatives enlarge and elongate. One of them 13) or superimposed (Fig. 6). Sometimes they becomes the ISTE mother cell which divides end blindly (Figs. 9, 12, 19), when they are unequally into two cells. The larger differentia­ away from main phloem strands. The number tes as an ISTE, the smaller one into a compan­ of ISTEs increases with distance from the vas­ ion cell (Figs. 29, 30). PAS positive spherical cular cambium. bodies appear in the developing ISTE (Fig. 32). ISTEs differentiate from the phloem ray pa­ The necrosis of the ISTE nucleus includes the renchyma cells in D. imperialis, G. angulosa change of a spherical nucleus to a spindle-shaped and V. volkameriaefolia and from interfascicu­ one (Figs. 28, 29), a marked decrease in stain­ lar parenchyma cells in Xanthium strumarium. ability of the nucIeus with Feulgen reagent and The salient developmental features have been traced in the former three taxa. The differen- (text continued on page 262) General explanation of Figures 1-32. All the seetions are of bark. Fig. 1-3, 28-32. Transverse sections. Fig. 14-17. Radial longitudinal sections. Fig. 4-9, 10-13, 18-20,21-27. Tangential longitudinal sections. Hollow arrow indicates isolated sieve tube element(s): ISTE(s); curved arrow indicates sieve plate and dart indicates companion cell. Fig. 1-9. I: Interfascicular region of Aristolochia indic4, x 158. - 2: Interfascicular region of Xan­ thium strumarium, x 349. - 3: Secondary phloem ray in Vernonia volkameriaefolia, x 502. - 4: Solitary ISTE in nonfunctional secondary phloem of V. volkameriaefolia, x 500. - 5: ISTE bridge in V. volkameriaefolia, x 500. - 6: Superimposed ISTEs in V. volkameriaefolia, x 500. - 7: Irregu­ lar arrangement of ISTE group in V. volkameriaefolia, x 500. - 8: ISTE at the corner of a ray in nonfunctional secondary phloem of V. volkameriaefolia, x 457. - 9: Blind ending of an ISTE in Dahlia imperialis, x 55. Fig. 10-20. 10: ISTE bridge at the corner of a ray in functional secondary phloem of Dahlia impe­ rialis, x 143. - 11: ISTE group in ray of functional secondary phloem of D. imperialis, x 46. - 12: ISTE group in nonfunctional secondary phloem of Strychnos nux-vomica, x 400. - 13: ISTE in nonfunctional secondary phloem of Gynura angulosa, x 414. - 14: ISTE cut transversely in Verno­ nia volkameriaefolia, x 580. - 15-17: Compound sieve plates of ISTE in V. volkameriaefolia (sa: sieve area, sp: sieve pore), 15, x 628; 16, x 549; 17, x 594. - 18-19: Blind endingofISTEin V. volkameriaefolia and D. imperialis; 18, x 914; 19, x 300. - 20: ISTE with L type of companion cell in D. imperialis, x 414. Fig.21-32. 21: Initials of isolated sieve tube element (zig-zag arrow) in Gynura angulosa, x 534. - 22: Differentiating ISTE in Dahlia imperialis, x 126. - 23: ISTE initial in D. imperialis, x 571. - 24-25: NucIear division of ISTE initial in D. imperialis (n: nucIeus); 24, x 526; 25, x 777. - 26: Nuclear division of ISTE initial in G. angulosa, x 457. - 27: Cytokinesis in ISTE initial of D. impe­ rialis, x 517. - 28: Arginine rich histone in ISTE initial and derivatives in D. imperialis, x 594. - 29: Necrosis of ISTE nucIeus (note change in shape) in D. imperialis (nn: necrotic nucIeus), x 722. - 30: EnucIeate ISTE having companion cell with DNA rich nucIeus in D. imperialis, x 830. - 31: Development of ISTE after two longitudinal divisions in Vernonia volkameriaefolia, x 342. - 32: PAS positive bodies (PB) in developing ISTE of D. imperialis, x 960. Downloaded from Brill.com09/29/2021 04:06:16AM via free access - 0\ <'l IAWA Bulletin n.s., Vol. 4 (4),198300 257 Downloaded from Brill.com09/29/2021 04:06:16AM via free access '" M 00 258 IAWA Bulletirt n.s., Vol. 4 (4),1983 Downloaded from Brill.com09/29/2021 04:06:16AM via free access 0- - <"> IAWA Bulletin n.s., Vol. 4 (4),1983co 259 Downloaded from Brill.com09/29/2021 04:06:16AM via free access t0 Table I. Dimensional details of isolated sieve tube elements. 0\ 0 Plant name STELin J..Ill1 STEWinJ..lll1 L/W No. of LSA / STE MINLSADinJ..lll1 MLSADinJ..lll1 SPPD inJ..lll1 LSAPDinJ..lll1 SPPD/ LSAPD Mean & SD Mean&SD Mean & SD Mean & SD Mean & SD Mean & SD Mean & SD Vemonia volkameriaefolia 87.4 ±31.2 26.9 ± 5.6 3.2 11.0 ± 5.8 1.4 ± 0.4 5.7 ± 2.3 1.4 ± 0.1 1.1 ± 0.2 1.2 Gynura angulosa 128.0 ± 51.0 21.0 ± 4.4 6.0 13.0 ± 11.8 1.2 ± 0.5 3.3 ± 1.7 1.5 ± 0.2 1.0 ± 0.1 1.3 Dahlia imperialis 48.8 ± 26.9 23.3 ± 3.3 2.0 4.0 ± 2.25 2.6 ± 1.5 5.5 ± 2.3 1.6 ± 0.3 1.1 ± 0.2 1.3 STEL = Sieve tube element length; STEW = Sieve tube element width; L/W = Sienderness ratio; LSA = Lateral sieve area; STE =Sieve tube element; MIN LSAD = Minimum diameter of lateral sieve area; MLSAD = Maximum diameter of lateral sieve area; SPPD = Sieve plate pore diameter; LSAPD = Lateral sieve area pore diameter; SD = Standard deviation.
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