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Clostridium Aerotolerans Sp INTERNATIONALJOURNAL OF SYSTEMATICBACTERIOLOGY, Apr. 1987, p. 102-105 Vol. 37, No. 2 0020-7713/87/020102-04$02.OO/O Copyright 0 1987, International Union of Microbiological Societies Clostridium aerotolerans sp. nov. a Xylanolytic Bacterium from Corn Stover and from the Rumina of Sheep Fed Corn Stover N. 0. VAN GYLSWYK" AND J. J. T. K. VAN DER TOORN Anaerobic Microbiology Division, Laboratory for Molecular and Cell Biology, Onderstepoort 01 10, South Africa Two strains of sporeforming, xylan-digesting, rod-shaped bacteria were isolated from a lo-* dilution of rumen ingesta taken from two sheep fed a corn stover ration. To determine the source of these strains, we examined the corn stover and isolated two strains of similar bacteria from it. The four strains were remarkably tolerant to oxygen although they did not grow in liquid medium that was shaken while exposed to air. They fermented a wide variety of carbon sources and produced formic, acetic, and lactic acids, ethanol, carbon dioxide, and hydrogen from xylan. The deoxyribonucleic acid base composition was 40 mol% guanine plus cytosine. The name proposed for these strains is Clostridiurn aerotolerans; the type strain is strain XSA62 (ATCC 43524). Two strains of sporeforming bacteria were found among a in which glucose (5 g/liter) replaced xylan. Spores were large number of bacteria isolated from colonies that pro- stained with malachite green (10). Motility was determined duced clearing zones in xylan (3%) agar medium inoculated after cells were grown overnight on maintenance slants. with a loh8 dilution of rumen ingesta from sheep fed corn Growth in basal medium containing cellobiose (5 g/liter) stover diets (14) as briefly described by van der Toorn and from which reducing agent (cysteine and sulfide) had been vim Gylswyk (15). To discover the origin of these bacteria, deleted was used to determine whether a low redox potential we examined the feed (corn stover) and isolated two strains was required. Tolerance to oxygen was assessed with 5-ml of bacteria with similar characteristics. In this paper, the amounts of basal medium in 50-ml Erlenmeyer flasks characteristics of the four strains are described; it is pro- (loosely stoppered with sterile aluminium caps), from which posed that they can be included in a new species, Clostrid- carbonate and reducing agent had been omitted. Resazurin iurn aerotolerans. (1 mg/liter) replaced indigo carmine (5 mg/liter) in the me- dium which contained clarified rumen fluid (40%), yeast MATERIALS AND METHODS extract (5 ghiter), and glucose (5 g/liter). For each 5 ml of medium, the inoculum was 0.5 ml of the water of syneresis Strains. Strains X8A62 and X4B63 were isolated from the from maintenance slants incubated overnight. In all cases, rumina of two different sheep adapted to corn stover fed ad inocula were examined microscopically for the presence of lib and supplemented with a protein-mineral mixture (14). an abundance of healthy cells. The mixture contained casein (46%), fish meal (41%), and Fermentation acids and alcohols were determined after minerals plus trace elements (12.5%). The corn stover diet growth in basal medium containing 20 g of xylan per liter. was fed (calculated on a daily basis) so that crude protein For acetylmethylcarbinol production, basal medium was formed 13.8% of total intake of solids. Strain X2 was isolated supplemented with cellobiose (5 g/liter) and glucose (10 from 2 g of corn stover (from the same batch as that fed to g/liter), and indigo carmine was deleted. the sheep) added to 200 ml of sterile, anaerobic diluent The medium used for the gelatin liquefaction test con- containing inorganic salts. This mixture was treated with a tained cellobiose (2 g/liter) and 120 g of gelatin per liter. Waring blender run for 3 min at half speed followed by Casein digestion was tested with a similar medium in which treatment with an Ultra-Turrax homogenizer (Janke and gelatin was replaced by 10 g of casein (Koch-Light Labora- Kunkel, Staufen i Br., Federal Republic of Germany) oper- tories Ltd., Colnbrook, England) per liter. Hydrogen sulfide ating at 20,000 rpm for 30 s. Strain X9 was isolated from a production was assessed in medium containing 5 g of separate sample of 4 g of stover treated in the same way. cellobiose and 10 g of peptone (tryptic digest of casein; E. Culture techniques and media. The techniques used and Merck AG, Darmstadt, Federal Republic of Germany) per the composition of basal medium and maintenance medium liter, and sulfide was deleted. Production of indole and were the same as those described before (16) except that a-methylindole was tested in medium containing 5 g of Na2C03(4 g/liter) replaced NaHC03 in these two media as cellobiose and 10 g of Casitone (Difco) per liter. Indigo well as in other media and that the maintenance medium carmine was deleted. For ability to reduce nitrate, cellobiose (used in preparing maintenance slants) contained 5 g of xylan (5 ghiter) and KN03 (1 g/liter, added in filter-sterilized (catalogue no. 95590; Fluka AG Buchs, Switzerland) per solution) were included in basal medium without a reducing liter. Soluble carbohydrate energy sources were included in agent and indigo carmine. Lipase activity was tested in basal sterile media in separate filter-sterilized or (for glucose) medium plus cellobiose (10 g/liter), agar (15 g/liter), and heat-sterilized solutions. tributyrin (Merck; ca. 10 g/liter). The presence or absence of The strains were isolated from basal medium (in roll catalase was assessed with maintenance slants containing 5 g bottles) containing xylan (30 g/liter) and agar (20 g/liter). of cellobiose per liter as the sole added energy source. For Colony morphology was examined after growth in this determining urease activity, cell suspensions were incubated medium but to which was added 5 g of yeast extract (Difco with basal medium to which was added filter-sterilized urea Laboratories, Detroit, Mich.) per liter and also in a medium solution to give a final concentration of 20 g of urea per liter. Basal medium containing 1.4 g of sodium DL-lactate served * Corresponding author. to test for ability to utilize lactate. 102 VOL. 37. 1987 CLOSTRIDIUM AEROTOLERANS SP. NOV. 103 Yeast extract (Difco) and peptone (Difco) at 5 g/liter each caps) free of added reducing agent but exposed to air. In this were incorporated in the following media: (i) medium to test medium growth is observed within 7 or 8 h after inoculation for ability to grow in the absence of carbohydrate in which and can be seen as a turbid layer on the bottom of the flasks clarified rumen fluid was used; and (ii) medium to test for where the medium is reduced with respect to resazurin. ability to grow in the absence of rumen fluid, carbonate, and About 15 min after gentle shaking of the flasks to suspend the C02 (C02 was replaced by N2 in the gas phase), which bacteria, the resazurin is almost completely reduced. On contained 5 g of cellobiose per liter. The effects of peptone standing, the upper layer of medium again becomes at least (10 g/liter) and yeast extract (5 g/liter) as growth stimulants partly oxidized with respect to resazurin, and after incuba- were examined with the inclusion of either or both in media tion for a further period of 16 to 17 h, growth appears as a containing 5 g of cellobiose per liter. For guanine-plus- dense layer on the bottom of the flasks with only the cytosine (G+C) determination, cells were grown in basal uppermost layer of medium showing some color due to medium supplemented with glucose and Difco yeast extract oxidized resazurin. At this stage vigorous shaking is required (5 g of each per liter). to dislodge the bacteria from the bottom of the flask where Media in which lowering of the pH served as a criterion of most growth has occurred. Incubation in a shaking water growth contained 0.1 X the usual concentration of carbonate, bath of inoculated medium prepared in the same way does and the gas phase was COrNTH* in a ratio of approxi- not produce any growth, and the medium is not reduced with mately 10:85:5. Also included were a medium for testing respect to resazurin even after 3 days. tolerance to different temperatures which contained Cells from 16-h cultures on agar slopes containing either cellobiose (5 g/liter), media for testing ability to ferment cellobiose or xylan are gram negative. Very little, if any, different carbohydrate energy sources (5 g of each per liter), stain is adsorbed if crystal violet is the only stain used, and a medium for assessing the types of gases produced Vegetative cells are straight rods often with somewhat during the fermentation of xylan (5 g/liter). In the last case, pointed ends (Fig. 1A and B). They usually occur singly. the CO2 content of the gas phase had to be low to facilitate Their width varies from 0.4 to 0.6 pm, and their length is the determination of C02produced during fermentation. The about 1.5 to 3.0 pm and sometimes more. Cells are motile by incubation temperature was 38 to 39°C except where indi- peritrichous flagella (Fig. 1B). Spores stain green with mal- cated, and the initial pH of the media varied from 6.6 to 6.9. achite green. They are terminal and distend the cells where Identification tests. Formic, acetic, D- and L-lactic and they are formed (Fig. 1C). They are slightly oval and succinic acids, and ethanol were determined by enzymatic measure about 0.6 by 0.8 pm.
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