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Transfer of Chainia Species to the Genus Streptomyces with Emended Description of Species
System. Appl. Microbiol. 8, 55-60 (1986)
Transfer of Chainia Species to the Genus Streptomyces with Emended Description of Species MICHAEL GOODFELLOW I , STANLEY T. WILLlAMS 2, and GRACE ALDERSON 3 1 2
3
Department of Microbiology, The Medical School, Framlington Place, Newcastle upon Tyne NE2 4HH, U.K. Department of Botany, University of Liverpool, Liverpool L69 3BX, U.K. School of Studies in Biomedical Sciences, University of Bradford, Bradford BD7 IDP, U.K.
Received August 12, 1985
Summary Chemical, genetical and phenetic data indicate a close relationship between the genus Chainia Thirumalachar and the genus Streptomyces Waksman and Henrici. It is proposed that the genus Chainia becomes a junior synonym of the genus Streptomyces and that the species it contains be named accordingly. Thus, it is proposed that Chainia antibiotica Thirumalachar, Chainia (lava Thirumalachar and Sukapure, Chainia fumigata Thirumalachar et aL, Chainia kunmingensis Ruan et aL, Chainia minutisclerotia Thirumalachar et al., Chainia nigra Thirumalachar, Chainia ochracea Kuznetsov, Chainia olivacea Thirumalachar and Sukapure, Chainia poonensis Thirumalachar, Chainia purpurogena Thirumalachar and Sukapure, Chainia rosea Thirumalachar, Chainia rubra Thirumalachar and Chainia violens Kalakoutskii and Krassilnikov be transferred to the genus Streptomyces as Streptomyces sclerotialus Pridham; Streptomyces (laviscleroticus (Pridham) nom. rev., Streptomyces fumigatiscleroticus (Pridham) nom. rev., Streptomyces kunmingensis (Ruan et al.) comb. nov., Streptomyces minutiscleroticus (Thirumalachar et al.) Pridham, Streptomyces niger (Thirumalacher) comb. nov., Streptomyces ochraceiscleroticus Pridham, Streptomyces olivaceiscleroticus Pridham, Streptomyces poonensis (Thirumalachar) Pridham, Streptomyces purpurogeneiscleroticus Pridham, Streptomyces roseiscleroticus Pridham, Streptomyces ruber (Thirumalachar) comb. nov. and Streptomyces violens (Pridham) nom. rev., respectively.
Key words: Chainia - Streptomyces - Taxonomy
Introduction
Thirumalachar (1955) introduced the genus Chainia for actinomycetes that form sclerotia either embedded in colonies growing on agar media or in shake flasks. Sclerotia, which may reach 100 ~m in diameter, consist of dense masses of enlarged, branching hyphae containing oil droplets (Sharples and Williams, 1976) cemented together by an amorphous material contammg L-2-3-diaminopropionic acid (Lechevalier et al., 1973). The type species, Chainia antibiotica, was described as lacking aerial mycelium but subsequently it was shown that representatives of this and other species assigned to the genus produced aerial hyphae and spore-chains very much like Streptomyces. The ability to form sclerotia is gradually lost on continued cultivation on laboratory media, while production of aerial mycelium is increased (Thirumalachar and Sukapure, 1964). Sclerotic granules have also been reported in
strains belonging to the genera Streptomyces (Lechevalier and Lechevalier, 1967; Williams et al., 1983; Goodfellow et al., 1985) and Streptoalloteichus (Tomita et al., 1978). The genus Chainia has been the subject of controversy with some workers considering that it should be regarded as a synonym of Streptomyces (Gattani, 1957; Bisset, 1959; Becker et al., 1965; Hutter, 1965; Lechevalier and Lechevalier, 1965, 1967; Prauser, 1970; Pridham and Tresner, 1974; Williams et aL, 1983; Goodfellow and Cross, 1984; Williams, 1985) and others giving reasons for the retention of the taxon (Lechevalier et aI., 1973; Cross and Goodfellow, 1973; Kutzner, 1981). Ten of the twelve species of Chainia included in the Approved Lists of Bacterial Names (Skerman et aL, 1980) had previously been assigned to the genus Streptomyces by Pridham (1970). Six of the ten species were described in the eighth
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M. Goodfellow, S. T. Williams, and G.Alderson
edition of Bergey's Manual of Determinative Bacteriology (Pridham and Tresner, 1974), the remaining four were cited as species incertae sedis (Table 1). Only the former, together with Streptomyces sclerotia Ius, were included in the Approved Lists of Bacterial Names (Skerman et al., 1980) as Streptomyces species. Recently, an additional Chainia species, Chainia kunmingensis was described (Ruan et al., 1985). Chainiae have DNA rich in guanine (G) plus cytosine (C) (Tsyganov et al., 1966, 1970; Yamaguichi, 1967), share a high DNA homology with Streptomyces (Yamaguichi, 1967; Kroppenstedt et al., 1981; Stackebrandt et al., 1981), are susceptible to polyvalent streptomycete phages (Prauser and Falta, 1968; Prauser, 1976; Kroppenstedt et al., 1981; Wellington and Williams, 1981), contain streptomycete-like fatty acids (Tsyganov et al., 1970), menaquinones (Collins et al., 1984; Alderson et al., 1985) and polar lipids (Lechevalier et al., 1977), cluster with Streptomyces species in numerical phenetic surveys (Williams et al., 1981, 1983) and have a wall chemotype I, that is, they contain LL-diaminopimelic acid and glycine but no characteristic sugars in the peptidoglycan (Pridham and Lyons, 1969; Lechevalier and Lechevalier, 1970; Schleifer and Kandler, 1972). Since these properties outweigh the transient ability of chainiae to produce sclerotia it is proposed that the genus Chainia (Thirumalachar, 1955) becomes a junior synonym of the genus Streptomyces (Waksman and Henrici, 1943) and that the species it contains be named accordingly. It is also proposed that Chainia antibiotica (Thirumalachar, 1955), Chainia (lava (Thirumalachar and Sukapure, 1964), Chainia fumigata (Thirumalachar et al., 1966), Chainia minutisclerotica (Thirumalachar et al., 1965), Chainia ochracea (Kuznetsov, 1962), Chainia olivacea (Thirumalachar and Sukapure, 1964), Chainia poonensis (Thirumalachar, 1960), Chainia purpurogena (Thirumalachar and Sukapure, 1964), Chainia rosea
(Thirumalachar, 1966) and Chainia vio/ens (Kalakoutskii and Krassilnikov, 1960) become synonyms of Streptomyces sclerotialus Pridham, 1970, Streptomyces (laviscleroticus (Pridham, 1970) nom. rev., Streptomyces fumigatiscleroticus (Pridham, 1970) nom. rev., Streptomyces minutiscleroticus (Thirumalachar et al., 1965) Pridham,
1970, Streptomyces ochraceiscleroticus (Pridham, 1970), Streptomyces olivaceiscleroticus (Pridham, 1970), Streptomyces poonensis (Thirumalachar, 1960) Pridham, 1970, Streptomyces purpurogeneiscleroticus (Pridham, 1970), Streptomyces roseiscleroticus (Pridham, 1970) and Streptomyces violens (Pridham, 1970) nom. rev., respectively. Emended descriptions are given for Streptomyces sclerotia Ius and for the three species for which names have been revived. Chainia kunmingensis (Ruan et al., 1985), Chainia nigra (Thirumalachar) Shirling and Gottlieb, 1972 and Chainia rubra (Thirumalachar) Shirling and Gottlieb, 1972 also have properties consistent with their classification in the genus Streptomyces (Shirling and Gottlieb, 1972; Lechevalier et al., 1977; Williams et ai, 1983; Collins et al., 1984; Ruan et al., 1985). Accordingly, it is proposed that Chainia kunmingensis, Chainia nigra and Chainia rubra be reclassified in the genus Streptomyces (Waksman and Henrici, 1943) as Streptomyces kunmingensis (Ruan et al.) comb. nov., Streptomyces niger (Thirumalachar) comb. nov., and Streptomyces ruber (Thirumalachar) comb. nov., respectively.
Descriptions Description of Streptomyces (laviscleroticus (Pridham) nom. rev.
(la. vi. scle. rot'i. cus. L. adj. (lavus, yellow; L. neut. n. sclerotium; M. L. adj. (laviscleroticus, yellow sclerotium, referring to yellow and ability to form sclerotia. The description is taken from information derived from
Table 1. Nomenclature and classification of Chainia species Skerman et al. 1980
Pridham 1970
Chainia antibiotica Thirumalachar 1955 Streptomyces sclerotia/us nom. nov. Chainia {lava Thirumalachar & Sukapure 1964Streptomyces {laviscleroticus nom. nov. Chainia fumigata Thirumalachar et al. 1966 Streptomyces fumigatiscleroticus nom. nov. Chainia minutisclerotica Thirumalachar et al. Streptomyces minutiscleroticus (Thirumalachar 1966 et al.) comb. nov. Chainia nigra Thirumalachar1955 Streptomyces ochraceiscleroticus nom. nov. Chainia ochracea Kuznetsov 1962 Streptomyces o/ivaceiscleroticus nom. nov. Chainia olivacea Thirumalachar & Sukapure 1964 Streptomyces poonensis (Thirumalachar) Chainia poonensis Thirumalachar 1960 comb. nov. Chainia purpurogena Thirumalachar & Suka- Streptomyces purpurogeniscleroticus nom. nov. pure 1964 Streptomyces roseiscleroticus nom. nov. Chainia rosea Thirumalachar 1966 Chainia rubra Thirumalachar 1955 Chainia violens Kalakoutskii and Krassilnikov Streptomyces violens (Kalakoutskii and Krassilnikov) comb. nov. 1960
• Cited as Species incertae sedis.
Pridham and Tresner 1974 ·Streptomyces sclerotia/us Pridham • Streptomyces {laviscleroticus Pridham • Streptomyces fumigatiscleroticus Pridham Streptomyces minutiscleroticus (Thirumalachar) Pridham Streptomyces ochraceiscleroticus Pridham Streptomyces o/ivaceiscleroticus Pridham Streptomyces poonensis (Thirumalachar) Pridham Streptomyces purpurogeneisclerosis Pridham Streptomyces roseiscleroticus Pridham
• Streptomyces vio/ens Pridham
Transfer of Chainia Species to the Genus Streptomyces
several sources (Thirmalachar and Sukapure, 1964; Shirling and Gottlieb, 1972; Williams et al., 1983; Collins et al., 1984; Ruan et al., 1985). Aerobic, Gram-positive, mesophilic organism with extensively branched substrate and aerial mycelium. Spiral spore chains of smooth surfaced spores borne on the aerial mycelium. Yellow brown substrate mycelium. Produces sclerotia. Does not form melanin pigments. Adenine, aesculin, casein, elastin, gelatin, guanine, hypoxanthine, starch and urea are degraded but allantoin, chitin, lecithin, pectin, testosterone, L-tyrosine, xanthine and xylan are not. Hydrogen sulphide is produced but nitrate is not reduced. L-arabinose, cellobiose, D-fructose, D-galactose, D-glucose, glycerol, D-Iactose, maltose, D-mannitol, D-mannose, raffinose, L-rhamnose, salicin, trehalose and D-xylose are used a sole carbon sources but adonitol, mesoinositol, inulin, D-melibiose, D-melezitose, sucrose and xylitol are not. Acid is formed from D-lactose, D-mannitol, a-methylD-glucoside, L-rhamnose, sucrose and D-xylose but not from adonitol, dulcitol, meso-erythritol, meso-inositol, Dmelibiose, raffinose and D-sorbitol. Grows on DL-a-amino-n-butyric acid, L-arginine, Lhistidine, potassium nitrate, L-serine, L-threonine and Lvaline, but not on L-cysteine, L-hydroxyproline, Lmethionine or L-phenylalanine as sole nitrogen source. Growth at 37 and 45°C but not at 10 0e. Tolerant to sodium azide (0.01 %, w/v) and sodium chloride (4%, w/v) but not to phenol (0.1 %, w/v). Sensitive to rifampicin and sodium chloride (7%, w/v). Does not show antimicrobial activity against Aspergillus niger LIV 131, Bacillus subtilis NCIB 3610, Candida albicans CBS 562, Escherichia coli NCIB 9132, Micrococcus luteus NCIB 196, Pseudomonas {luorescens NCIB 9046, Saccharomyces cerevisiae CBS 1171 or Streptomyces murinus ISP 5091. The organism contains major amounts of octahydrogenated menaquinones with nine isoprene units. Isolated from soil, Pimpri, India. Type strain: ATCC 19347 (= DSM 43152 = IFO 12998 = ISP 5270 = KCC 3100 = NCIB 11008 = RIA 833). Description of Streptomyces (Pridham) nom. rev.
fumigatiscleroticus
fu. mi. ga'ti. scle. rot'i. cus. L. masc. n. fumus, smoke, steam; L. suf. atus, provided with; L. part. adj. fumigatus, smoked; L. neut. n. sclerotium; M. L. adj. fumigatiscleroticus, smoked sclerotium, referring to smoke colour and ability to form sclerotia. The description is taken from information derived from Thirumalachar et al. (1966) and Ruan et al. (1985). Aerobic, Gram-positive organism with extensively branched substrate and aerial mycelium. Produces spiral spore chains and shows brownish-black crusty growth. Produces sclerotia. Melanin pigments are not formed. Degrades gelatin, starch, urea and xanthine. Nitrate is reduced but hydrogen sulphide is not produced.
57
L-arabinose, glucose, lactose, maltose, D-mannitol, mannose, sorbitol, sucrose and D-xylose are used as sole carbon sources but dulcitol, galactose, glycerol, inulin, meso-inositol, L-rhamnose, salicin and sucrose are not. Acid is produced from D-lactose, D-mannitol, Dmelibiose, a-methyl-D-glucoside, raffinose, L-rhamnose, D-sorbitol and D-xylose but not from adonitol, dulcitol, meso-erythritol, meso-inositol and sucrose. Growth at 42°C but not at 10°e. Shows antimicrobial activity against Bacillus subtilis, Escherichia coli and Staphylococcus aureus but not against Candida albicans, Cryptococcus neoformans or
Saccharomyces cerevisiae.
Isolated from soil, Pimpri, India. Type strain: ATCC 19345 (= DSM 43154 = IFO 12999 = KCC 3101 = RIA 884). Description of Streptomyces kunmingensis (Ruan et al.)
comb. nov.
kun. mingo en'sis. M. L. adj. kunmingensis, pertaining to Kunming, a province of South China. The description is taken from Ruan et al. (1985). Aerobic, Gram-positive organism with extensively branched substrate and aerial mycelium. Chains of conidia in loose spirals are occasionally borne on the aerial hyphae. Colonies usually yellowish tan to dark orange yellow, depending on the medium, with or without yellowish white aerial mycelium. Produces sclerotia (5 to 25 !!m) in and on the surface of agar media. Melanin pigments not formed. Adenine, aesculin, casein, hypoxanthine, starch, tyrosine and xanthine are degraded but gelatin and urea are not. Phosphatase and nitrate reductase are produced. L-arabinose, D-mannitol, raffinose, L-rhamnose and Dxylose are used as sole carbon sources but meso-inositol and sucrose are not. Acid is formed from L-arabinose, cellobiose, fructose, galactose, glucose, glycerol, meso-inositol, D-lactose, maltose, D-mai:mitol, mannose, melibiose, a-methyl-Dglucoside, a-methyl-D-xyloside, raffinose, L-rhamnose, salicin, sucrose, trehalose and D-xylose but not from adonitol, dulcitol, meso-erythritol or D-sorbitol. Growth at 10 and 37"C but not at 42°e. The wall peptidoglycan contains LL-diaminopimelic acid as the major diamino acid. The organism has a type II phospholipid pattern (sensu Lechevalier et al., 1977). The mole % G plus C content of the DNA of the type strain is 71.3%. Isolated from soil at Dagyanlow's Lake in Kunming, Peoples' Republic od China. Type strain ATCC 35682. Description of Streptomyces niger (Thirumalachar) comb. nov. ni'ger. L. adj. niger, black. The description is taken from information derived from several sources (Thirumalachar, 1955; Shirling and Gottlieb, 1972; Williams et al., 1983; Ruan et al., 1985). Aerobic, Gram-positive, mesophilic organism with extensively branched substrate and aerial mycelium. Spiral
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M. Goodfellow, S. T. Williams, and G.Alderson
spore chains of smooth surfaced spores borne on the aerial mycelium. The aerial spore mass is white and the substrate mycelium green. A green soluble pigment is produced. Sclerotia are formed in 7 to 14 days on agar media. Melanin pigments are not produced. Adenine, aesculin, allantoin, arbutin, casein, elastin, guanine, hypoxanthine, pectin, starch, testosterone, L-tyrosine, xanthine and xylan are degraded but chitin, lecithin and urea are not. Nitrate is reduced but hydrogen sulphide is not produced. Adonitol, L-arabinose, cellobiose, D-fructose, D-galactose, D-glucose, meso-inositol, inulin, D-Iactose, D-mannitol, D-mannose, D-melezitose, D-melibiose, raffinose, Lrhamnose, salicin, sucrose, trehalose, xylitol and D-xylose are all used as sole cabon sources. Acid is formed from adonitol, meso-erythritol, mesoinositol, D-Iactose, D-mannitol, D-melibiose, a-methyl-Dglucoside, raffinose, L-rhamnose, sucrose and D-xylose but not from dulcitol or D-sorbitol. Grows on DL-a-amino-butyric acid, L-arginine, L-histidine, L-phenylalanine, potassium nitrate, L-serine, Lthreonine and L-valine, but not on L-cysteine or Lmethionine as sole nitrogen source. Growth at 10, 37 and 45°C but not at 4°e. Tolerant to phenol (0.1%, w/v), sodium azide (0.02%, w/v) and sodium chloride (10%, NaCl). Resistant to rifampicin but not to sodium chloride (13%, w/v). Antibiotic activity shown against Bacillus subtilis NCIB 3610 and Saccharomyces cerevisiae CBS 1171 but not against Aspergillus niger LIV 131, Candida albicans CBS 562, Escherichia coli NCIB 9132, Micrococcus luteus NCIB 196, Pseudomonas f1uorescens NCIB 9046 or Streptomyces murinus ISP 5091. The organism has a type II phospholipid pattern (sensu Lechevalier et al., 1977) and contains major amounts of hexahydrogenated menaquinones with nine isoprene units (Collins et al., 1984). Type strain ATCC 17756 (= DSM 43049 = IFO 13362 = ISP 5302 = KCC 3158 = RIA 1323). Description of Streptomyces ruber (Thirumalachar) comb. nov. rub' er. L. adj. ruber, red. The description is taken from information derived from
Thirumalachar (1955), Shirting and Gottlieb (1972) and Ruan et al. (1985).
Aerobic, Gram-positive, mesophilic organism with extensively branched substrate and aerial mycelium. Spiral spore chains containing 10 to over 50 spores are borne on the aerial mycelium. The spore surface is smooth or bears very short spines. The aerial spore mass is white or red and the substrate mycelium red. Sclerotia are formed in 14 to 21 days on agar media. Melanin pigments are not produced. Xanthine is degraded but urea is not. Nitrate reductase is not formed. L-arabinose, D-fructose, D-glucose, D-mannitol, rhamnose and D-xylose are used as sole carbon sources but
raffinose is not utilised. Acid is formed from D-Iactose, D-mannitol, D-
melibiose, a-methyl-D-glucoside, L-rhamnose, sucrose and D-xylose but not from adonitol, dulcitol, meso-erythritol, meso-inositol, raffinose, and D-sorbitol. Growth at 42°C but not at 100e. The organism contains major amounts of hexa- and octa-hydrogenated menaquinones with nine isoprene units (Collins et al., 1984). Type strain: ATCC 17754 (= ISP 5304 = KCC 3131 = NCIB 10983). Emendation of Streptomyces sclerotialus Pridham.
scle. rot. i' a. Ius. L.neut. n. sclerotium, sclerotium; M. L. adj. sclerotialus, pertaining to sclerotia. The description is taken from information derived from several sources (Thirumalachar, 1955; Thirumalachar and Sukapure, 1964; Williams et aI, 1983; Ruan et al., 1985). Aerobic, Gram-positive, mesophilic organism with extensively branched substrate and aerial mycelium. Spiral spore chains of smooth surfaced spores borne on the aerial mycelium. Colonies yellow-brown with a white aerial spore mass. Sclerotia formed. Melanin pigments not produced. Adenine, aesculin, allantoin, arbutin, casein, chitin, elastin, gelatin, guanine, hypoxanthine, starch, testosterone, L-tyrosine, urea and xanthine are degraded but lecithin, pectin and xylan are not. Hydrogen sulphide is produced but nitrate is not reduced. Adonitol, L-arabinose, cellobiose, D-fructose, D-galactose, D-glucose, meso-inositol, inulin, D-lactose, D-mannitol, D-mannose, D-melezitose, D-melibiose, raffinose, Lrhamnose, salicin, sucrose, trehalose, xylitol and D-xylose are all used as sole carbon sources. Acid is formed from adonitol, meso-erythritol, mesoinositol, D-lactose, D-mannitol, D-melibiose, a-methyl-Dglucoside, raffinose, L-rhamnose, sucrose and D-xylose but not from dulcitol or D-sorbitol. Grows on DL-a-amino-n-butyric acid, L-arginine, Lhistidine, L-phenylalanine, potassium nitrate, L-serine, Lthreonine and L-valine, but not on L-cysteine, L-hydroxyproline or L-methionine as sole nitrogen source. Growth at 37 and 45°C but not at lO°e. Tolerant to phenol (0.1 %, w/v), sodium azide (0.01 %, w/v) and sodium chloride (10%, w/v). Sensitive to rifampicin and sodium chloride (13%, w/v). Does not show antimicrobial activity against Aspergillus niger LIV 131, Bacillus subtilis NCIB 3610, Candida albicans CBS 562, Escherichia coli NCIB 9132, Micrococcus luteus NCIB 196, Pseudomonas f1uorescens NCIB 9046, Saccharomyces cerevisiae CBS 1171 or Streptomyces murinus ISP 5091. The peptidoglycan contains LL-diaminopimelic acid
(Becker et al., 1965; Pridham and Lyons, 1969) and is of the A3y type (Stackebrandt et al., 1981). The organism also contains major amounts of octahydrogenated menaquinones with nine isoprene units (Alderson et al., 1985). Isolated from soil, Poona, India. Type strain: ATCC 15721 (= DSM 43032 = IFO 12246 = ISP 5269 = KCC 3039 = RIA 1317).
Transfer of Chainia Species to the Genus Streptomyces
Description of Streptomyces violens (Pridham) comb. nov.
vi. o. lens. L. adj. violens, raging but probably from L. fern. n. viola, violet, referring to pink to violet pigment produced by the organism on some media. The description is taken from information derived from several sources (Kalakoutskii and Krassilinikov, 1960; Shirling and Gottlieb, 1972; Williams et aI., 1983; Ruan et al., 1985). Aerobic, Gram-positive, mesophilic organism with extensively branched substrate and aerial mycelium. Spiral spore chains of smooth surfaced spores produced. Produces a yellow brown substrate mycelium, the pigment is pH sensitive. Forms sclerotia. Melanin pigments are not produced but forms a yellow brown diffusible pigment. Adenine, aesculin, allantoin, arbutin, casein, elastin, gelatin, guanine, hypoxanthine, pectin, starch, testosterone, L-tyrosine, urea, xanthine and xylan are degraded but lecithin and chitin are not. Hydrogen sulphide is produced but nitrate is not reduced. Adonitol, L-arabinose, cellobiose, D-fructose, D-galactose, D-glucose, meso-inositol, inulin, D-Iactose, D-mannitol, D-mannose, raffinose, L-rhamnose, salicin, trehalose, xylitol and D-xylose are used as sole carbon sources but sucrose is not. Acid is formed from adonitol, meso-erythritol, mesoinositol, D-Iactose, D-mannitol, D-melibiose, a-methyl-Dglucoside, raffinose, L-rhamnose, D-sorbitol, sucrose and D-xylose but not from dulcitol. Grows on DL-a-amino-n-butyric acid, L-arginine, Lhistidine, potassium nitrate, L-serine and L-valine, but not from L-cysteine, L-hydroxyproline, L-methionine or Lphenylalanine. Growth at 10, 37 and 45 °C but not at 4°C. Tolerant to phenol (0.01 %, w/v), sodium azide (0.01 %, w/v) and sodium chloride (10%, w/v) but sensitive to rifampicin and sodium chloride (13 %, w/v). Shows antimicrobial activity against Micrococcus luteus NCIB 196 but not against Aspergillus niger LIV 131, Bacillus subtilis NCIB 3610, Candida albicans CBS 562, Escherichia coli NCIB 9132, Pseudomonas fluorescens NCIB 9046, Saccharomyces cerevisiae CBS 1171 or Streptomyces murinus ISP 5091. The peptidoglycan contains LL-diaminopimelic acid (Becker et aI., 1965). The mole % G plus C of the DNA of the type strains has been reported to be 70.9 (Tsyganov et aI., 1966) and 71.5 (Yamaguichi, 1967). Isolated from soil, Armenia, USSR. Type strain: ATCC 15898 (= IFO 12557 = ISP 5597 = KCC 3072 = RIA 565).
References Alderson, G., Goodfellow, M. , Minnikin, D. E.: Menaquinone composition in the classification of Streptomyces and other sporoactinomycetes. ]. gen. Microbiol. 131 , 1671-1679 (1985)
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Becker, B., Lechevalier, M. P., Lechevalier, H. A.: Chemical composition of cell wall preparations from strains of various form-genera of aerobic actinomycetes. Appl. Microbiol. 13, 236-243 (1965) Bisset, K. A.: The morphology and natural relationships of saprophytic actinomycetes. Progr. Industr. Microbiol. 1, 29-43 (1959) Collins, M. D., Faulkner, M. , Keddie, R. M.: Menaquinone composition of some sporeforming actinomycetes. System. Appl. Microbiol. 5, 20-29 (1984) Cross, T., Goodfellow, M. : Taxonomy and classification of the actinomycetes. In: Actinomycetales: Characteristics and Practical Importance (G. Sykes and F. A. Skinner, eds.), pp. 11-112. London, Academic Press 1973 Gattani, M. L.: Production of sclerotic granules by Streptomyces spp. Nature (Lond.) 180, 1293-1294 (1957) Goodfellow, M., Cross, T.: Classification. In: The Biology of the Actinomycetales (M . Goodfellow, M. Mordarski and S. T. Williams, eds), pp. 7-164. London, Academic Press 1984 Goodfellow, M., Williams, S. T., Alderson, G.: Transfer of Actinosporangium violaceum Krasil'nikov and Yuan, Actinosporangium vitaminophilum Shomura et al. and Actinopycnidium caeruleum Krasil'nikov to the genus Streptomyces, with amended descriptions of the species. System. App\. Microbiol. 8, 61-64 (1986) Hutter, R.: Die Gattungen der Actinomycetales. Path. Microbiol. 28, 567-574 (1965) Kalakoutskii, L. V., Krassilnikov, N. A.: Formation of sclerotia in actinomycetes and the sytematic position of genus Chainia. In: Biology of antibiotic-producing actinomycetes (Rautenshtein, Ya, I., ed.). Akad. Nauk. SSSR Inst. Mikrobiol. Trudy 8, 45-55 (1960) Kroppenstedt, R. M., Korn-Wendisch, F., Fowler, V. j., Stackebrandt, E.: Biochemical and molecular genetic evidence for a transfer of Actinoplanes armeniacus into the family Steptomycetaceae. Zbl. Bakt. Hyg., I. Abt. Orig. C 2, 254-262 (1981 ) Kutzner, H. J.: Streptomycetaceae. In: The Prokaryotes: A Handbook of Habitats, Isolation and Identification of Bacteria (M. P. Starr, H. Stolp, H. G., Truper, A. Balows and H. G. Schlegel, eds.), pp. 2028-2090. Berlin, Springer-Verlag 1981 Kuznetsov, V. D.: A new species of Chainia (English translation). Microbiology 31, 435-439 (1962) Lechevalier, H. A., Lechevalier, M. P.: Classification des actinomycetes aerobic basees sur leur morphologie et leur composition chimique. Ann. Inst. Pasteur (Paris) 108, 662-673 (1965) Lechevalier, H. A., Lechevalier, M. P.: Biology of actinomycetes. Ann. Rev. Microbiol. 21, 71-100 (1967) Lechevalier, M. P., Lechevalier, H. A.: Composition of wholecell hydrolysates as a criterion in the classification of aerobic actinomycetes. In: The Actinomycetales (H. Prauser, ed.), pp. 311-316. lena, Gustav Fischer Verlag 1970 Lechevalier, M. P., De Bievre, c., Lechevalier, H.: Chemotaxonomy of aerobic actinomycetes: phospholipid composition. Biochem. System. Ecol. 5, 249-260 (1977) Lechevalier, M. P., Lechevalier, H. A., Heintz, C. E.: Morphological and chemical nature of the sclerotia of Chainia olivacea Thirumalachar and Sukapure of the order Actinomycetales. Int. J. system. Bact. 23, 157-170 (1973) Prauser, H. : Characters and genera arrangement in the Actinomycetales. In: The Actinomycetales (H. Prauser, ed.), pp. 407-418. lena, Gustav Fischer Verlag 1970 Prauser, H.: Host-phage relationships in nocardioform organisms. In: The Biology of the Nocardiae (M. Goodfellow, G. H. Borwnell and J. A. Serrano, eds.), pp. 266-284. London, Academic Press 1976
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Prauser, H., Falta, R.: Phagensensibilitiit, Zellwand-Zusammensetzung und Taxonomie von Actinomyceten. Z. allg. Mikrobio!. 8, 39-46 (1968) Pridham, T. G.: New names and new combinations in the order Actinomycetales Buchanan 1917. U.S. Dept. Agr. Techn. Bull. 1424, 1-55 (1970) Pridham, T. G., Lyons, jr., A. J.: Progress in the clarification of the taxonomic and nomenclatural status of some problem actinomycetes. Dev. indo Microbiol. 10, 183-221 (1969) Pridham , T. G., Tresner, H. D.: Genus I. Streptomyces Waksman and Henrici 1943,339. In: Bergey's Manual of Determinative Bacteriology, eighth Edition (R . E. Buchanan and N. E. Gibbons, eds.), pp. 747-829. Baltimore, Williams and Wilkins 1974 Ruan, }.-S., Lechevalier, M. P., Jiang, c.-L., Lechevalier, H. A.: Chainia kunmingensis, a new actinomycete species found in soil. Int. J. system. Bact. 35, 164-168 (1985) Schleifer, K. H., Kandler, 0.: Peptidoglycan types of bacterial cell walls and their taxonomic implications. Bact. Rev. 36, 407-477 (1972) Sharples, G. P., Williams, S. T. : Development and fine structure of sclerotia and spores of the actinomycete Chainia oliuacea. Microbios 15, 37-47 (1976) Shirling, E. B., Gottlieb, D.: Comparative description of type strains of Streptomyces. V. Additional descriptions. Int. J. system. Bact. 22, 265-394 (1972) Skerman, V. B. D., McGowan, V. , Sneath, P. H. A.: Approved lists of bacterial names. Int. J. system Bact. 30, 225-420 (1980)
Stackebrandt, E., Wunner-Fussl, B., Fowler, V.}., Schleifer, K.H. : Deoxyribonucleic acid homologies and ribosomal ribonucleic acid similarities among sporeforming members of the order Actinomycetales. Int. J. system. Bact. 31, 420-431 (1981) Thirumalachar, M. }.: Chainia, a new genus of the Actinomycetales. Nature (Lond.) 176, 934-935 (1955) Thirumalachar, M. }.: See Kalakoutskii, L. V., Krassilnikov, N. A., in: Biology of antibiotic-producing actinomycetes (Rautenshtein, Ya.I., ed.). Akad. Nauk. SSSR Inst. Mikrobiol. T rudy 8, 45-55 (1960) Thirumalachar, M. }., Sukapure, R. S.: Studies on species of the
genus Chainia from India. Hindustan Antibiot. Bull. 6, 157-166 (1964) Thirumalachar, M.}., Rahalkar, P. W., Deshmukh, P. V., Sukapure, R. S.: Production of aburamycin by Chainia minutisclerotica, a new species of Actinomycetes. Hindustan Antibiot. Bull. 8, 6-9 (1965)
Thirmualachar, M. }., Sukapure, R. S., Rahalkar, P. W., Gopalkrishnan, K. S.: Studies on species of the genus Chainia. India II. Hindustan Antibiot. Bull. 9, 10-14 (1966) Thirumalachar, M. }., Sukapure, R. S., Rahalkar, P. W., Gopalkrishnan, K. S.: Studies on species of the genus Chainia from India. II. Hindustan Antibiot. Bull. 9, 10-15 (1966)
Tomita, K., Uenoyama, Y., Numata, K.-I., Sasahira, T., Hoshino, Y., Fujisawa, K.-I., Tsukiura, H., Kawaguichi, H.: Streptoalloteichus, a new genus of the Actinoplanaceae. J. Antibiot. 31, 497-570 (1978)
Tsyganou, V. A., Namestnikova, V. P., Krassikova, M. V.: Composition of DNA in members of different genera of actinomycetes. Mikrobilogiya 35, 75-78, (1966) (in Russian)
Tsyganov, V. A., Efimoua, T. P., Zhukova, R. A., Koneu, Yu. E.: New forms of actinomycetes and taxonomic significance of some of their metabolites. In : The Actinomycetales (H. Prauser, ed.), pp. 329-335. Jena, Gustav Fischer Verlag 1970 Waksman, S. A., Henrici, A. T.: the nomenclature and classification of the actinomycetes. J. Bact. 46, 337-341 (1943) Wellington, E. M. H., Williams, S. T. : Host ranges of phage isolated to Streptomyces and other genera. Zbl. Bakt., Suppl. 11,93-98 (1981) Williams, S. T.: The taxonomy of streptomycetes - problems and perspectives. Front. Appl. Microbiol. 1, in press Williams, S. T., Wellington, E. M. H., Goodfellow, M., Alderson, G., Sackin, M., Sneath, P. H. A.: The genus Streptomyces - a taxonomic enigma. Zbl. Bakt., Suppl. 11, 47-57 (1981) Williams, S. T., Goodfellow, M., Alderson, G., Wellington, E. M. H., Sneath, P. H. A., Sack in, M. }.: Numerical classification of Streptomyces and related genera. J. gen. Microbiol. 129, 1743-1813 (1983) Yamaguichi, T.: Similarity in DNA of various morphologically distinct actinomycetes. J. gen. app!. Microbiol. 13, 63-71 (1967)
Dr. M. Goodfellow, Dept. of Microbiology, The Medical School, Framlington Place, Newcastle upon Tyne NE2 4HH, U.K.
Transfer of Chainia Species to the Genus Streptomyces with Emended Description of Species MICHAEL GOODFELLOW I , STANLEY T. WILLlAMS 2, and GRACE ALDERSON 3 1 2
3
Department of Microbiology, The Medical School, Framlington Place, Newcastle upon Tyne NE2 4HH, U.K. Department of Botany, University of Liverpool, Liverpool L69 3BX, U.K. School of Studies in Biomedical Sciences, University of Bradford, Bradford BD7 IDP, U.K.
Received August 12, 1985
Summary Chemical, genetical and phenetic data indicate a close relationship between the genus Chainia Thirumalachar and the genus Streptomyces Waksman and Henrici. It is proposed that the genus Chainia becomes a junior synonym of the genus Streptomyces and that the species it contains be named accordingly. Thus, it is proposed that Chainia antibiotica Thirumalachar, Chainia (lava Thirumalachar and Sukapure, Chainia fumigata Thirumalachar et aL, Chainia kunmingensis Ruan et aL, Chainia minutisclerotia Thirumalachar et al., Chainia nigra Thirumalachar, Chainia ochracea Kuznetsov, Chainia olivacea Thirumalachar and Sukapure, Chainia poonensis Thirumalachar, Chainia purpurogena Thirumalachar and Sukapure, Chainia rosea Thirumalachar, Chainia rubra Thirumalachar and Chainia violens Kalakoutskii and Krassilnikov be transferred to the genus Streptomyces as Streptomyces sclerotialus Pridham; Streptomyces (laviscleroticus (Pridham) nom. rev., Streptomyces fumigatiscleroticus (Pridham) nom. rev., Streptomyces kunmingensis (Ruan et al.) comb. nov., Streptomyces minutiscleroticus (Thirumalachar et al.) Pridham, Streptomyces niger (Thirumalacher) comb. nov., Streptomyces ochraceiscleroticus Pridham, Streptomyces olivaceiscleroticus Pridham, Streptomyces poonensis (Thirumalachar) Pridham, Streptomyces purpurogeneiscleroticus Pridham, Streptomyces roseiscleroticus Pridham, Streptomyces ruber (Thirumalachar) comb. nov. and Streptomyces violens (Pridham) nom. rev., respectively.
Key words: Chainia - Streptomyces - Taxonomy
Introduction
Thirumalachar (1955) introduced the genus Chainia for actinomycetes that form sclerotia either embedded in colonies growing on agar media or in shake flasks. Sclerotia, which may reach 100 ~m in diameter, consist of dense masses of enlarged, branching hyphae containing oil droplets (Sharples and Williams, 1976) cemented together by an amorphous material contammg L-2-3-diaminopropionic acid (Lechevalier et al., 1973). The type species, Chainia antibiotica, was described as lacking aerial mycelium but subsequently it was shown that representatives of this and other species assigned to the genus produced aerial hyphae and spore-chains very much like Streptomyces. The ability to form sclerotia is gradually lost on continued cultivation on laboratory media, while production of aerial mycelium is increased (Thirumalachar and Sukapure, 1964). Sclerotic granules have also been reported in
strains belonging to the genera Streptomyces (Lechevalier and Lechevalier, 1967; Williams et al., 1983; Goodfellow et al., 1985) and Streptoalloteichus (Tomita et al., 1978). The genus Chainia has been the subject of controversy with some workers considering that it should be regarded as a synonym of Streptomyces (Gattani, 1957; Bisset, 1959; Becker et al., 1965; Hutter, 1965; Lechevalier and Lechevalier, 1965, 1967; Prauser, 1970; Pridham and Tresner, 1974; Williams et aL, 1983; Goodfellow and Cross, 1984; Williams, 1985) and others giving reasons for the retention of the taxon (Lechevalier et aI., 1973; Cross and Goodfellow, 1973; Kutzner, 1981). Ten of the twelve species of Chainia included in the Approved Lists of Bacterial Names (Skerman et aL, 1980) had previously been assigned to the genus Streptomyces by Pridham (1970). Six of the ten species were described in the eighth
56
M. Goodfellow, S. T. Williams, and G.Alderson
edition of Bergey's Manual of Determinative Bacteriology (Pridham and Tresner, 1974), the remaining four were cited as species incertae sedis (Table 1). Only the former, together with Streptomyces sclerotia Ius, were included in the Approved Lists of Bacterial Names (Skerman et al., 1980) as Streptomyces species. Recently, an additional Chainia species, Chainia kunmingensis was described (Ruan et al., 1985). Chainiae have DNA rich in guanine (G) plus cytosine (C) (Tsyganov et al., 1966, 1970; Yamaguichi, 1967), share a high DNA homology with Streptomyces (Yamaguichi, 1967; Kroppenstedt et al., 1981; Stackebrandt et al., 1981), are susceptible to polyvalent streptomycete phages (Prauser and Falta, 1968; Prauser, 1976; Kroppenstedt et al., 1981; Wellington and Williams, 1981), contain streptomycete-like fatty acids (Tsyganov et al., 1970), menaquinones (Collins et al., 1984; Alderson et al., 1985) and polar lipids (Lechevalier et al., 1977), cluster with Streptomyces species in numerical phenetic surveys (Williams et al., 1981, 1983) and have a wall chemotype I, that is, they contain LL-diaminopimelic acid and glycine but no characteristic sugars in the peptidoglycan (Pridham and Lyons, 1969; Lechevalier and Lechevalier, 1970; Schleifer and Kandler, 1972). Since these properties outweigh the transient ability of chainiae to produce sclerotia it is proposed that the genus Chainia (Thirumalachar, 1955) becomes a junior synonym of the genus Streptomyces (Waksman and Henrici, 1943) and that the species it contains be named accordingly. It is also proposed that Chainia antibiotica (Thirumalachar, 1955), Chainia (lava (Thirumalachar and Sukapure, 1964), Chainia fumigata (Thirumalachar et al., 1966), Chainia minutisclerotica (Thirumalachar et al., 1965), Chainia ochracea (Kuznetsov, 1962), Chainia olivacea (Thirumalachar and Sukapure, 1964), Chainia poonensis (Thirumalachar, 1960), Chainia purpurogena (Thirumalachar and Sukapure, 1964), Chainia rosea
(Thirumalachar, 1966) and Chainia vio/ens (Kalakoutskii and Krassilnikov, 1960) become synonyms of Streptomyces sclerotialus Pridham, 1970, Streptomyces (laviscleroticus (Pridham, 1970) nom. rev., Streptomyces fumigatiscleroticus (Pridham, 1970) nom. rev., Streptomyces minutiscleroticus (Thirumalachar et al., 1965) Pridham,
1970, Streptomyces ochraceiscleroticus (Pridham, 1970), Streptomyces olivaceiscleroticus (Pridham, 1970), Streptomyces poonensis (Thirumalachar, 1960) Pridham, 1970, Streptomyces purpurogeneiscleroticus (Pridham, 1970), Streptomyces roseiscleroticus (Pridham, 1970) and Streptomyces violens (Pridham, 1970) nom. rev., respectively. Emended descriptions are given for Streptomyces sclerotia Ius and for the three species for which names have been revived. Chainia kunmingensis (Ruan et al., 1985), Chainia nigra (Thirumalachar) Shirling and Gottlieb, 1972 and Chainia rubra (Thirumalachar) Shirling and Gottlieb, 1972 also have properties consistent with their classification in the genus Streptomyces (Shirling and Gottlieb, 1972; Lechevalier et al., 1977; Williams et ai, 1983; Collins et al., 1984; Ruan et al., 1985). Accordingly, it is proposed that Chainia kunmingensis, Chainia nigra and Chainia rubra be reclassified in the genus Streptomyces (Waksman and Henrici, 1943) as Streptomyces kunmingensis (Ruan et al.) comb. nov., Streptomyces niger (Thirumalachar) comb. nov., and Streptomyces ruber (Thirumalachar) comb. nov., respectively.
Descriptions Description of Streptomyces (laviscleroticus (Pridham) nom. rev.
(la. vi. scle. rot'i. cus. L. adj. (lavus, yellow; L. neut. n. sclerotium; M. L. adj. (laviscleroticus, yellow sclerotium, referring to yellow and ability to form sclerotia. The description is taken from information derived from
Table 1. Nomenclature and classification of Chainia species Skerman et al. 1980
Pridham 1970
Chainia antibiotica Thirumalachar 1955 Streptomyces sclerotia/us nom. nov. Chainia {lava Thirumalachar & Sukapure 1964Streptomyces {laviscleroticus nom. nov. Chainia fumigata Thirumalachar et al. 1966 Streptomyces fumigatiscleroticus nom. nov. Chainia minutisclerotica Thirumalachar et al. Streptomyces minutiscleroticus (Thirumalachar 1966 et al.) comb. nov. Chainia nigra Thirumalachar1955 Streptomyces ochraceiscleroticus nom. nov. Chainia ochracea Kuznetsov 1962 Streptomyces o/ivaceiscleroticus nom. nov. Chainia olivacea Thirumalachar & Sukapure 1964 Streptomyces poonensis (Thirumalachar) Chainia poonensis Thirumalachar 1960 comb. nov. Chainia purpurogena Thirumalachar & Suka- Streptomyces purpurogeniscleroticus nom. nov. pure 1964 Streptomyces roseiscleroticus nom. nov. Chainia rosea Thirumalachar 1966 Chainia rubra Thirumalachar 1955 Chainia violens Kalakoutskii and Krassilnikov Streptomyces violens (Kalakoutskii and Krassilnikov) comb. nov. 1960
• Cited as Species incertae sedis.
Pridham and Tresner 1974 ·Streptomyces sclerotia/us Pridham • Streptomyces {laviscleroticus Pridham • Streptomyces fumigatiscleroticus Pridham Streptomyces minutiscleroticus (Thirumalachar) Pridham Streptomyces ochraceiscleroticus Pridham Streptomyces o/ivaceiscleroticus Pridham Streptomyces poonensis (Thirumalachar) Pridham Streptomyces purpurogeneisclerosis Pridham Streptomyces roseiscleroticus Pridham
• Streptomyces vio/ens Pridham
Transfer of Chainia Species to the Genus Streptomyces
several sources (Thirmalachar and Sukapure, 1964; Shirling and Gottlieb, 1972; Williams et al., 1983; Collins et al., 1984; Ruan et al., 1985). Aerobic, Gram-positive, mesophilic organism with extensively branched substrate and aerial mycelium. Spiral spore chains of smooth surfaced spores borne on the aerial mycelium. Yellow brown substrate mycelium. Produces sclerotia. Does not form melanin pigments. Adenine, aesculin, casein, elastin, gelatin, guanine, hypoxanthine, starch and urea are degraded but allantoin, chitin, lecithin, pectin, testosterone, L-tyrosine, xanthine and xylan are not. Hydrogen sulphide is produced but nitrate is not reduced. L-arabinose, cellobiose, D-fructose, D-galactose, D-glucose, glycerol, D-Iactose, maltose, D-mannitol, D-mannose, raffinose, L-rhamnose, salicin, trehalose and D-xylose are used a sole carbon sources but adonitol, mesoinositol, inulin, D-melibiose, D-melezitose, sucrose and xylitol are not. Acid is formed from D-lactose, D-mannitol, a-methylD-glucoside, L-rhamnose, sucrose and D-xylose but not from adonitol, dulcitol, meso-erythritol, meso-inositol, Dmelibiose, raffinose and D-sorbitol. Grows on DL-a-amino-n-butyric acid, L-arginine, Lhistidine, potassium nitrate, L-serine, L-threonine and Lvaline, but not on L-cysteine, L-hydroxyproline, Lmethionine or L-phenylalanine as sole nitrogen source. Growth at 37 and 45°C but not at 10 0e. Tolerant to sodium azide (0.01 %, w/v) and sodium chloride (4%, w/v) but not to phenol (0.1 %, w/v). Sensitive to rifampicin and sodium chloride (7%, w/v). Does not show antimicrobial activity against Aspergillus niger LIV 131, Bacillus subtilis NCIB 3610, Candida albicans CBS 562, Escherichia coli NCIB 9132, Micrococcus luteus NCIB 196, Pseudomonas {luorescens NCIB 9046, Saccharomyces cerevisiae CBS 1171 or Streptomyces murinus ISP 5091. The organism contains major amounts of octahydrogenated menaquinones with nine isoprene units. Isolated from soil, Pimpri, India. Type strain: ATCC 19347 (= DSM 43152 = IFO 12998 = ISP 5270 = KCC 3100 = NCIB 11008 = RIA 833). Description of Streptomyces (Pridham) nom. rev.
fumigatiscleroticus
fu. mi. ga'ti. scle. rot'i. cus. L. masc. n. fumus, smoke, steam; L. suf. atus, provided with; L. part. adj. fumigatus, smoked; L. neut. n. sclerotium; M. L. adj. fumigatiscleroticus, smoked sclerotium, referring to smoke colour and ability to form sclerotia. The description is taken from information derived from Thirumalachar et al. (1966) and Ruan et al. (1985). Aerobic, Gram-positive organism with extensively branched substrate and aerial mycelium. Produces spiral spore chains and shows brownish-black crusty growth. Produces sclerotia. Melanin pigments are not formed. Degrades gelatin, starch, urea and xanthine. Nitrate is reduced but hydrogen sulphide is not produced.
57
L-arabinose, glucose, lactose, maltose, D-mannitol, mannose, sorbitol, sucrose and D-xylose are used as sole carbon sources but dulcitol, galactose, glycerol, inulin, meso-inositol, L-rhamnose, salicin and sucrose are not. Acid is produced from D-lactose, D-mannitol, Dmelibiose, a-methyl-D-glucoside, raffinose, L-rhamnose, D-sorbitol and D-xylose but not from adonitol, dulcitol, meso-erythritol, meso-inositol and sucrose. Growth at 42°C but not at 10°e. Shows antimicrobial activity against Bacillus subtilis, Escherichia coli and Staphylococcus aureus but not against Candida albicans, Cryptococcus neoformans or
Saccharomyces cerevisiae.
Isolated from soil, Pimpri, India. Type strain: ATCC 19345 (= DSM 43154 = IFO 12999 = KCC 3101 = RIA 884). Description of Streptomyces kunmingensis (Ruan et al.)
comb. nov.
kun. mingo en'sis. M. L. adj. kunmingensis, pertaining to Kunming, a province of South China. The description is taken from Ruan et al. (1985). Aerobic, Gram-positive organism with extensively branched substrate and aerial mycelium. Chains of conidia in loose spirals are occasionally borne on the aerial hyphae. Colonies usually yellowish tan to dark orange yellow, depending on the medium, with or without yellowish white aerial mycelium. Produces sclerotia (5 to 25 !!m) in and on the surface of agar media. Melanin pigments not formed. Adenine, aesculin, casein, hypoxanthine, starch, tyrosine and xanthine are degraded but gelatin and urea are not. Phosphatase and nitrate reductase are produced. L-arabinose, D-mannitol, raffinose, L-rhamnose and Dxylose are used as sole carbon sources but meso-inositol and sucrose are not. Acid is formed from L-arabinose, cellobiose, fructose, galactose, glucose, glycerol, meso-inositol, D-lactose, maltose, D-mai:mitol, mannose, melibiose, a-methyl-Dglucoside, a-methyl-D-xyloside, raffinose, L-rhamnose, salicin, sucrose, trehalose and D-xylose but not from adonitol, dulcitol, meso-erythritol or D-sorbitol. Growth at 10 and 37"C but not at 42°e. The wall peptidoglycan contains LL-diaminopimelic acid as the major diamino acid. The organism has a type II phospholipid pattern (sensu Lechevalier et al., 1977). The mole % G plus C content of the DNA of the type strain is 71.3%. Isolated from soil at Dagyanlow's Lake in Kunming, Peoples' Republic od China. Type strain ATCC 35682. Description of Streptomyces niger (Thirumalachar) comb. nov. ni'ger. L. adj. niger, black. The description is taken from information derived from several sources (Thirumalachar, 1955; Shirling and Gottlieb, 1972; Williams et al., 1983; Ruan et al., 1985). Aerobic, Gram-positive, mesophilic organism with extensively branched substrate and aerial mycelium. Spiral
58
M. Goodfellow, S. T. Williams, and G.Alderson
spore chains of smooth surfaced spores borne on the aerial mycelium. The aerial spore mass is white and the substrate mycelium green. A green soluble pigment is produced. Sclerotia are formed in 7 to 14 days on agar media. Melanin pigments are not produced. Adenine, aesculin, allantoin, arbutin, casein, elastin, guanine, hypoxanthine, pectin, starch, testosterone, L-tyrosine, xanthine and xylan are degraded but chitin, lecithin and urea are not. Nitrate is reduced but hydrogen sulphide is not produced. Adonitol, L-arabinose, cellobiose, D-fructose, D-galactose, D-glucose, meso-inositol, inulin, D-Iactose, D-mannitol, D-mannose, D-melezitose, D-melibiose, raffinose, Lrhamnose, salicin, sucrose, trehalose, xylitol and D-xylose are all used as sole cabon sources. Acid is formed from adonitol, meso-erythritol, mesoinositol, D-Iactose, D-mannitol, D-melibiose, a-methyl-Dglucoside, raffinose, L-rhamnose, sucrose and D-xylose but not from dulcitol or D-sorbitol. Grows on DL-a-amino-butyric acid, L-arginine, L-histidine, L-phenylalanine, potassium nitrate, L-serine, Lthreonine and L-valine, but not on L-cysteine or Lmethionine as sole nitrogen source. Growth at 10, 37 and 45°C but not at 4°e. Tolerant to phenol (0.1%, w/v), sodium azide (0.02%, w/v) and sodium chloride (10%, NaCl). Resistant to rifampicin but not to sodium chloride (13%, w/v). Antibiotic activity shown against Bacillus subtilis NCIB 3610 and Saccharomyces cerevisiae CBS 1171 but not against Aspergillus niger LIV 131, Candida albicans CBS 562, Escherichia coli NCIB 9132, Micrococcus luteus NCIB 196, Pseudomonas f1uorescens NCIB 9046 or Streptomyces murinus ISP 5091. The organism has a type II phospholipid pattern (sensu Lechevalier et al., 1977) and contains major amounts of hexahydrogenated menaquinones with nine isoprene units (Collins et al., 1984). Type strain ATCC 17756 (= DSM 43049 = IFO 13362 = ISP 5302 = KCC 3158 = RIA 1323). Description of Streptomyces ruber (Thirumalachar) comb. nov. rub' er. L. adj. ruber, red. The description is taken from information derived from
Thirumalachar (1955), Shirting and Gottlieb (1972) and Ruan et al. (1985).
Aerobic, Gram-positive, mesophilic organism with extensively branched substrate and aerial mycelium. Spiral spore chains containing 10 to over 50 spores are borne on the aerial mycelium. The spore surface is smooth or bears very short spines. The aerial spore mass is white or red and the substrate mycelium red. Sclerotia are formed in 14 to 21 days on agar media. Melanin pigments are not produced. Xanthine is degraded but urea is not. Nitrate reductase is not formed. L-arabinose, D-fructose, D-glucose, D-mannitol, rhamnose and D-xylose are used as sole carbon sources but
raffinose is not utilised. Acid is formed from D-Iactose, D-mannitol, D-
melibiose, a-methyl-D-glucoside, L-rhamnose, sucrose and D-xylose but not from adonitol, dulcitol, meso-erythritol, meso-inositol, raffinose, and D-sorbitol. Growth at 42°C but not at 100e. The organism contains major amounts of hexa- and octa-hydrogenated menaquinones with nine isoprene units (Collins et al., 1984). Type strain: ATCC 17754 (= ISP 5304 = KCC 3131 = NCIB 10983). Emendation of Streptomyces sclerotialus Pridham.
scle. rot. i' a. Ius. L.neut. n. sclerotium, sclerotium; M. L. adj. sclerotialus, pertaining to sclerotia. The description is taken from information derived from several sources (Thirumalachar, 1955; Thirumalachar and Sukapure, 1964; Williams et aI, 1983; Ruan et al., 1985). Aerobic, Gram-positive, mesophilic organism with extensively branched substrate and aerial mycelium. Spiral spore chains of smooth surfaced spores borne on the aerial mycelium. Colonies yellow-brown with a white aerial spore mass. Sclerotia formed. Melanin pigments not produced. Adenine, aesculin, allantoin, arbutin, casein, chitin, elastin, gelatin, guanine, hypoxanthine, starch, testosterone, L-tyrosine, urea and xanthine are degraded but lecithin, pectin and xylan are not. Hydrogen sulphide is produced but nitrate is not reduced. Adonitol, L-arabinose, cellobiose, D-fructose, D-galactose, D-glucose, meso-inositol, inulin, D-lactose, D-mannitol, D-mannose, D-melezitose, D-melibiose, raffinose, Lrhamnose, salicin, sucrose, trehalose, xylitol and D-xylose are all used as sole carbon sources. Acid is formed from adonitol, meso-erythritol, mesoinositol, D-lactose, D-mannitol, D-melibiose, a-methyl-Dglucoside, raffinose, L-rhamnose, sucrose and D-xylose but not from dulcitol or D-sorbitol. Grows on DL-a-amino-n-butyric acid, L-arginine, Lhistidine, L-phenylalanine, potassium nitrate, L-serine, Lthreonine and L-valine, but not on L-cysteine, L-hydroxyproline or L-methionine as sole nitrogen source. Growth at 37 and 45°C but not at lO°e. Tolerant to phenol (0.1 %, w/v), sodium azide (0.01 %, w/v) and sodium chloride (10%, w/v). Sensitive to rifampicin and sodium chloride (13%, w/v). Does not show antimicrobial activity against Aspergillus niger LIV 131, Bacillus subtilis NCIB 3610, Candida albicans CBS 562, Escherichia coli NCIB 9132, Micrococcus luteus NCIB 196, Pseudomonas f1uorescens NCIB 9046, Saccharomyces cerevisiae CBS 1171 or Streptomyces murinus ISP 5091. The peptidoglycan contains LL-diaminopimelic acid
(Becker et al., 1965; Pridham and Lyons, 1969) and is of the A3y type (Stackebrandt et al., 1981). The organism also contains major amounts of octahydrogenated menaquinones with nine isoprene units (Alderson et al., 1985). Isolated from soil, Poona, India. Type strain: ATCC 15721 (= DSM 43032 = IFO 12246 = ISP 5269 = KCC 3039 = RIA 1317).
Transfer of Chainia Species to the Genus Streptomyces
Description of Streptomyces violens (Pridham) comb. nov.
vi. o. lens. L. adj. violens, raging but probably from L. fern. n. viola, violet, referring to pink to violet pigment produced by the organism on some media. The description is taken from information derived from several sources (Kalakoutskii and Krassilinikov, 1960; Shirling and Gottlieb, 1972; Williams et aI., 1983; Ruan et al., 1985). Aerobic, Gram-positive, mesophilic organism with extensively branched substrate and aerial mycelium. Spiral spore chains of smooth surfaced spores produced. Produces a yellow brown substrate mycelium, the pigment is pH sensitive. Forms sclerotia. Melanin pigments are not produced but forms a yellow brown diffusible pigment. Adenine, aesculin, allantoin, arbutin, casein, elastin, gelatin, guanine, hypoxanthine, pectin, starch, testosterone, L-tyrosine, urea, xanthine and xylan are degraded but lecithin and chitin are not. Hydrogen sulphide is produced but nitrate is not reduced. Adonitol, L-arabinose, cellobiose, D-fructose, D-galactose, D-glucose, meso-inositol, inulin, D-Iactose, D-mannitol, D-mannose, raffinose, L-rhamnose, salicin, trehalose, xylitol and D-xylose are used as sole carbon sources but sucrose is not. Acid is formed from adonitol, meso-erythritol, mesoinositol, D-Iactose, D-mannitol, D-melibiose, a-methyl-Dglucoside, raffinose, L-rhamnose, D-sorbitol, sucrose and D-xylose but not from dulcitol. Grows on DL-a-amino-n-butyric acid, L-arginine, Lhistidine, potassium nitrate, L-serine and L-valine, but not from L-cysteine, L-hydroxyproline, L-methionine or Lphenylalanine. Growth at 10, 37 and 45 °C but not at 4°C. Tolerant to phenol (0.01 %, w/v), sodium azide (0.01 %, w/v) and sodium chloride (10%, w/v) but sensitive to rifampicin and sodium chloride (13 %, w/v). Shows antimicrobial activity against Micrococcus luteus NCIB 196 but not against Aspergillus niger LIV 131, Bacillus subtilis NCIB 3610, Candida albicans CBS 562, Escherichia coli NCIB 9132, Pseudomonas fluorescens NCIB 9046, Saccharomyces cerevisiae CBS 1171 or Streptomyces murinus ISP 5091. The peptidoglycan contains LL-diaminopimelic acid (Becker et aI., 1965). The mole % G plus C of the DNA of the type strains has been reported to be 70.9 (Tsyganov et aI., 1966) and 71.5 (Yamaguichi, 1967). Isolated from soil, Armenia, USSR. Type strain: ATCC 15898 (= IFO 12557 = ISP 5597 = KCC 3072 = RIA 565).
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M. Goodfellow, S. T. Williams, and G.Alderson
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Dr. M. Goodfellow, Dept. of Microbiology, The Medical School, Framlington Place, Newcastle upon Tyne NE2 4HH, U.K.