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Chemical Nature of Major Cell Wall Constituents of Vaucheria and Dichotomosiphon with Special Reference to Their Phylogenetic Positions

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Chemical Nature of Major Cell Wall Constituents of Vaucheria and Dichotomosiphon with Special Reference to Their Phylogenetic Positions Bot• Mag. Tokyo 79: 634-643 (Hattori Commemoration Number, 1966) Chemical Nature of Major Cell Wall Constituents of Vaucheria and Dichotomosiphon with Special Reference to their Phylogenetic Positions by Masaakira MAEDA*,Kazuko KURODA**, Yoshihiko IRIKI** *, Mitsuo CHIHARA*** )~, Kazutosi NIsIzAWA** and Tomoo MIWA** ReceivedOctober 13, 1966 For the elucidation of phylogenetic relations of plants, morphological characters as well as the mode of life cycle, especially in lower plants, have thus far been considered essential. Besides these, the chemical nature of the substances constitut- ing the plant body or produced therein, though invisible directly, has recently been regarded as important and used in taxonomy and phylogeny. The chemical nature of substances used for plant taxonomy is highly diversified and includes flavonoids, terpenes, fatty acids, hydrocarbons and others. Concerning these situations the book of Swains' may be consulted. In Japan the works of Kariyone2' on biflavones and leaf waxes of conifers, and those of Fu jita3>on essential oils are worth noting. Recent advances in biochemical genetics indicate that the chemical nature of plant products is controlled by genes in the same sense as are morphological characters, thus form- ing the basis for a chemical taxonomy. As for cell wall constituents, F. von Wettstein4' stated earlier that the presence or absence of chitin is important as a criterion for the classification of fungi. In the domain of algae, Miwa et al.5,6,7,studied the chemical nature of some siphonous green algae in relation to their phylogenetic positions. They found that Siphonales can be divided sharply into two groups with respect to the nature of the polysac- charide which, in place of cellulose, constitutes the main cell wall substance and is obtainable in form of crude fiber. One group, including the genera, Codium and Derbesia, is characterized by the presence of mannan ; and the other group, to which Caulerpa, Bryopsis, Pseudodichotomosiphon,Halimeda, Udotea, Chlorodesmisand Avrainvillea belong, contains xylan in its place. Such a grouping is in accord with the classification by Feldmanng' of Siphonales based chiefly on the morphological characters, except for the position of Bryopsis. In the present study the chemical nature of the main cell wall material of Vaucheria and Dichotomosiphonwas investigated with a view to obtaining informa- tion concerning the phylogenetic position of these algae. The systematic position of Vaucheria in the system of algae seems not to be completely settled. Some phycologists have placed the Vaucheriaceae in the order Siphonales (Oltmanns 19229',Printz 1927'0', Okamura 193011',193612', Ijengar 1951'3', Fritsch 1935'40, and later Fritsch (1954)15'has proposed to establish an independent * Departmentof Biochemistry, Faculty of Engineeringand Science,Saitama University, Urawa,Japan. ** BotanicalInstitute , Faculty of Science,Tokyo Kyoiku University,Tokyo, Japan. ***Biological Laboratory , Departmentof Education,Shinshu University, Nagano, Japan. ****National Science Museum , Ueno,Tokyo, Japan. Oct.-Nov., 1966 MAEDA, M., et al. 635 order Vaucheriales in the siphonous green algae. Contrary to these views other workers would remove the Vaucheriaceae from Chlorophyta and place them in the Xanthophyceae (Bohlin 190116, Feldmann 19458', Smith 195017), Papenfuss 1955180. As stated above, the main cell wall material of the siphonous green algae con sists either of xylan or of mannan and not cellulose. As to the cell wall constituents of members of the Xanthophyceae no definite information is available at present Cellulose has been considered to be generally lacking in this group of algae, though. its occurrence was reported in Tribonema as a result of X-ray study19). In Vaucheria the chemical nature of the main cell wall constituent has not beena thoroughly clarified. Thus, Nicolai and Prestonl9) concluded on the basis of physical investigation and staining reactions, that the skeletal substance of the cell wall of Vaucheria is not cellulose, however, others have reported that cellulose is present. (e. g., Printz 192710), Fritsch 195514) but Chadefaud (1945)20) separated this genus fromm the Vaucheriaceae and placed it in a new family Dichotomosiphonaceae. Since the fact that the main cell wall constituent of Dichotomosiphon was callose but not cel- lulose was considered as one of the most important properties distinguishing thiss genus from Vaucheria, the present study was undertaken to obtain some information about the systematic relation, if any, between Vaucheria and Dichotomosiphon by comparing chemical nature of main cell wall contituent. Materials and Methods Vaucheria : Two species were used. V. sessilis was harvested at the end of March in a shallow stream near Tokyo ; V. longicaulis, a marine species, was obtainedd from Okinawa. Dichotomosiphon : D. tuberoses was collected in Okinawa. All of the materials were carefully freed from attached impurities, washedd thoroughly with fresh water, air-dried and finally dried at 105° to constant weight and ground to coarse powder. A part of fresh-water Vaucheria was preserved in ethanol for use in histochemical tests. Ash.-The sample was heated at about 550 ° in an electric muffle furnace for 12 hr. The residue was dissolved in 6 N nitric acid and the insoluble matter filtered off. The filtrate was evaporated and dried to constant weight. Nitrogen.-Total nitrogen was determined by micro-Kjeldahl method. Pentosan and methylpentosan.-These were estimated by the method of Kruger- Tollens-Krober2l) and Elett-Tollens-Mayer22), respectively. Reducing sugars.--In most cases Somogyi's method23) and for pentose Tracey's method24' were applied. In the presence of xylose, glucose was estimated by the use of glucose oxidase, the enzyme preparation employed (" Deoxin ", Nagase Co., Ltd.) had almost negligible action upon xylose. Paper chromatography.-Ascending chromatography was carried out on Toya filter paper No. 52 with n-butanol-pyridine-water (6: 4 : 3, v/v) as developer. For the detection of sugars, either acetone-silver nitrate or benzidine-trichloroacetic acid was sprayed. Infrared spectra.-A Nihon Bunko, type iR-S instrument was used. Measure- ment was made with the sample in a potassium bromide disk. X636 Bot. Mag. Tokyo Vol. 79 Results Vaucheria sessilis and V. longicaulis. Histochemical tests.-Staining reactions for cellulose were carried out with the fronds either preserved in alcohol or dried in air. Both the freshwater and marine species gave similar results. The cell wall stained pale blue with iodine and satu- rated zinc chloride and blue with iodine and concentrated sulfuric acid. Upon pre- treatment of the fronds with dilute hypochlorite solution, the color intensity increased significantly with both reagents. Iodine and concentrated calcium chloride stained the cell wall red. Analysis of whole fronds.-The ash, nitrogen, pentosan and methylpentosan contents of dried fronds were determined. The results are presented in Table 1. Table 1. Analysis of whole frond (Numerals on dry weight basis). Preparation of crude fiber.-Samples of both species were extracted respectively twice with hot 80% ethanol each time for 1 hr., then treated with 1% hydrochloric acid at 18-20° overnight. After washing with water the residue was heated with 1.25% sodium hydroxide at 100° for 30 min. The residue was bleached with dilute sodium hypochlorite (1 available chlorine) at room temperature for 1 hr. and freed from excess oxidant by washing with a dilute solution of sodium sulfite. The product was then heated with 1.25% sulfuric acid at 100° for 30 min. and washed thoroughly with water until free of sulfuric acid. Yield, 23% of dried frond. The crude fiber thus obtained showed, when examined under the microscope, an almost unchanged structure. The previously mentioned staining reactions for cellulose were clearer with those crude fiber preparation than with the original fronds. The fiber preparations showed distinct double refraction. Acid hydrolysis of crude fiber.-100 mg of each crude fiber preparation was mixed with 2 ml of 72% sulfuric acid and kept at 0° for 24 hr. The dark brown product obtained was diluted with water to 40 ml and heated at 100° for 2 hr. The hydro- lyzate was neutralized with barium carbonate and filtered. The filtrate was concen- trated in vacuo to a syrup which was then dissolved in hot ethanol. After filtration the ethanolic solution was evaporated and the product finally dissolved in 3 ml water. Paper chromatography showed the presence of glucose only. Enzymatic hydrolysis of crude fiber.-A commercial praparation of" Trichoderma- Cellulase " (Mei ji-Seika Co., Ltd.) was dissolved in water, dialyzed free of reducing substances and lyophilized. Each of the crude Vaucheria fiber preparations was ground to fine powder and 50 mg thereof was suspended in 1.5 ml of 0.1 M acetate buffer, pH 5.2, and incubated with 0.5 ml of 3% cellulase solution at 30° . After 24 hr. the reaction mixture was centrifuged and the clear supernatant fluid was examined chromatographically. Besides the major spot of glucose , several smaller spots of lower migration rates were found. The latter seemed to be oligosaccharides com- posed of glucose, since the extracts from two of these spots yielded upon acid hyro- lysis a single spot corresponding to glucose. Oct.-Nov., 1966 MAEDA, M., e t al. 637 Purification of crude fiber.-A sample of the crude fiber from V. sessilis was mixed well with ten times its weight of 17.5% sodium hydroxide; after 30 min. the mixture was diluted with an equal volume of water and centrifuged. The residue was washed successively with dilute acetic acid and with water, and dried. Yield 98.9%. This preparation showed also marked staining reactions for cellulose as well as distinct double refraction. Moreover, a sample was subjected to acetolysis25', and the product obtained was purified by repeated recrystallization from a mixture of chloro- form and methanol. Its melting point, 222°, and infrared spectrum agreed well with those of a-octaacetylcellobiose. Dichotomosiphon tuberoses. Analysis of whole fronds.-Quantitative determination of some of the constitu- ents are shown in Table 2.
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