Cooperative Description of Type Cultures of STREPTOMYCES III. Additional Species Descriptions From First and Second Studies

INTERNATIONAL JOURNAL O F SYSTEMATIC BACTERIOLOGY October 1968 Vol. 18, No.
4 Copyright 1968, Iowa State University P r e s s
pp. 279- 392
COOPERATIVE DESCRIPTION O F T Y P E CULTURES O F STREPTOMYCES III. ADDITIONAL SPECIES DESCRIPTIONS FROM FIRST AND SECOND STUDIES** Elwood B. Shirling and David Gottlieb Department of Botany and Bacteriology Ohio Wesleyan University, Delaware, Ohio, and Department of Plant Pathology, University of Illinois, Urbana, Illinois
ABSTRACT. M o r e t h a n 4 0 c o o p e r a t i n g l a b o r a t o r i e s r e p r e s e n t i n g 18 n a t i o n s a r e j o i n e d i n a n i n t e r n a t i o n a l e f fort to assemble and redescribe authentic type s t r a i n s o r acceptable neotype strains for the named species in the genera Streptomyces and Streptoverticillium. Part I1 of t h i s s e r i e s i n c l u d e d i l I h s t r a t e d e m e n d a t i o n s t o t h e d e s c r i p t i o n s f o r 1 0 0 t y p e s t r a i n s . P a r t I11 e x t e n d s the report t o include type s t r a i n s for a n additional 100 Streptomyces o r Streptoverticillium species. Each new d e s c r i ption i s based upon data submitted by t h r e e c o operating laboratories, using the standardized criteria and methods officially adopted for this purpose. Emendations for additional species will be included in r e ports to follow. Type .specimens f o r all redescribed species a r e deposited in the American Type Culture C o l l e c t i o n , T h e C e n t r a a l b u r e a u v o o r S c h i m m e l c u l t u r e s, T h e USSR R e s e a r c h I n s t i t u t e f o r A n t i b i o t i c s , a n d i n Japan.
- - - - - - - - - - - -
Collaborators and authors of descriDtions: Group A- 2* C r o s s , T. and A. Maciver. Postgraduate School of Studies i n Biological Sciences, Univ. of Bradford, Bradford 7 / Yorkshire, England. Johanides, Vera and T e a Blazekovic. L a b o r a t o r i j z a industrijsku mikrobiologiju, Tehnoloxki fakultet, Pierottijeva 6, Zagreb, Jug0 slavija. T r e j o , W. E. R. Squibb and Sons, New Brunswick, New J e r s e y , U. S. A. A letter-number combination is a s signed t o e a c h collaborating group. T o conserve space, t h i s code is used t o indicate the author group for e a c h emended description. The l e t t e r A indicates collaborating groups f o r t h e first 130 type s t r a i n s distributed for study; B designates groups participating in the second study. Reprints of t h i s a r t i c l e m a y be purchased f r o m the International Micr.obiologica1 Fund, 22 1 Science Hall, Iowa State University, Ames, Iowa 50010 U. S. A. P r i c e f o r single copies: 562.00.
*:
**
280
INTERNATIONAL
JOURNAL
Group A-3 Crook, Kenneth E., J r . and C a r o l S. C a s s i d y . B r i s t o l L a b s . , Inc. S y r a c u s e , New York, U . S. A. d e V r i e s , G. A. C e n t r a a l b u r e a u voor S c h i m m e l c u l t u r e s , B a a r n , Netherlands. Krasil'nikov, N. A. Inst. Microbiology. Academy of S c i e n c e s , Moscow B- 133, U . S. S. R. G r o u p A-8 O l i v e r , T. J. Abbott L a b o r a t o r i e s , N . Chicago, Illinois, U. S. A. Spyvee, J. and J. Elliott, Boots P u r e Drug C o . , Ltd., Antibiot. and F e r m . Div., Nottingham, England. Nakazawa, K. , M. Shibata, K. Yamamoto, E. Higashide, T . Iwasa and T . Hasagawa. Takeda C h e m . Ind., Ltd., Juso-Higashiyodogawaku, O s a k a , Japan. G r o u p A-9 N i s h i m u r a , H., M. Mayama and K. T a w a r a . Shionogi R e s . Lab., Shionogi and Co., Ltd., F u k u s h i m a - k u , O s a k a , Japan. P r i d h a m , T. G. N o r t h e r n Utilization R e s . and Dev. Div., P e o r i a , Illinois, U. S. A. Wognicka, Wanda. PaGstwowy Zakaad Higieny. W a r s zawa 12, ul. C h o c i m s k a 24, Poland. Group A- 12 Margalith, P. L a b o r . of M i c r o b i o l . , Dept. of Food and Biotechnology, Technion, Haifa, I s r a e l . Vernon, T . R. Dept. Scientific and Ind. R e s . , P r i v a t e Bag, Auckland C1, New Z e a l a n d . Williams, S . T . H a r t l e y Botan. L a b o r . , Univ. of Liverpool, Liverpool, England. G r o u p A- 1 3 k e h a r e k , Z . and Alena kigicovg. Czech. Acad. of Science, Inst. of Microbiol. , P r a g u e 4, Budejovicka No. 1083, Czechoslovakia. S a n c h e z - M a r r o q u i n , A. M i a m i 40, Mexico. 18 / D. F., Mexico. Williams, S. T. H a r t l e y Botan. L a b o r . , Univ. of Liverpool, Liverpool, England. G r o u p B-1 S a n c h e z - M a r r o q u i n , A. M i a m i 40, Mexico 18 / D . F . , Mexico. Baldacci, E . , G. F a r i n a and R. Locci. 1st. Patol. Vegetale, Univ. Milano, Milano, Via C e l o r i a N. 2, Milano, Italy, Gauze, G . F . , T . P . P r e o b r a z h e n s k a y a , E.S. Kudrina, T.S. Maximova and M.A. Sveshnikova. Acad. Med. S c i e n c e s of U . S. S. R . , Inst. New Antibiotics, Bolshaia P i r o g o v s k a i a 11, Moscow, U. S. S. R.
SYSTEMATIC
BACTERIOLOGY
28 1
GrouD B- 2 Anderson, Lucia E. and S a r a Wold. P a r k e , Davis and Co. , Detroit, Michigan, U. S. A. C r o s s , T. and A. Maciver. Postgraduate School of Studies in Biological Sciences, Univ. of Bradford, Bradford, 7 , Yorkshire, England. Johanides, Vera and T e a Blazekovic. Laboratorij za h d u s t r i j s k u mikrobiologiju, Tehnolo2ki fakultet, Pierottijeva 6, Zagreb, Jugoslavija. Group B-3 Radziminska, h a . Hoffman- LaRoche, Nutley, New J e r s e y , U. S. A. d e v r i e s , G.A. Centraalbureau voor Schimmelcultures, Baarn, Nether lands. Krasil'nikov, N. A. Inst. Microbiology. Academy of Sciences, Moscow B- 133, U. S. S. R.
Group B-4
Crook, Kenneth E., Jr. and C a r o l S. Cassidy. B r i s t o l Labs., Inc., Syracuse, New York, U. S . A. Mach, F. Inst. fur Mikrobiologie, Greifswald, Ludwig- Jahn-Str. 15, G e r m a n Democratic Republic Kuznetsov, V. D. USSR R e s e a r c h Inst. f o r Antibiotics, Moscow, U. S. S. R.
Group B-5 Oliver, T. J. Abbott Laboratories, N. Chicago, Illinois, U. S.A. P r a u s e r , H. Inst. Mikrobiol. Exptl. Therapie, Jena, Beuthenbergstr. 11, G e r m a n Democratic Republic. Szab6, I. Magyar Tudomgnyos Akadgmia, Talajtana ks Agrokgmiai Kutat6 Intgzete, Budapest, Hungary. Group B-6 kehagek, Z . and Alena &i&vocz. Czech. Acad. of Science, Inst. of Microbiol., P r a g u e 4, Budejovicka No. 1083, Cz.echoslovakia. Mitchell, T . G. T o r r y R e s e a r c h Station, Aberdeen Scotland. Tsyganov, V. A. Res. Inst. of Antibiotics, Leningrad, L-20, U. S . S . R. Group B-8 McClung, Norvel M.* and Gene Michaels. Dept. Microbiol.. Univ. Georgia, Athens, G a . , U. S.A. * ( P r e s e n t a d d r e s s : Dept. Bot. and Bact., Univ. South Florida, Tampa, Florida. ) Spyvee, J. and J. Elliott. Boots P u r e D r u g Co., Ltd., Antibiot. and Ferm. Div., Nottingham, England: Nakazawa, K. , M. Shibata, K. Yamamoto, E. Higashide, T. Iwasa and T . Hasegawa. Takeda Chem. Ind., Ltd., Juso-Higashiyodogawaku, Osaka, Japan.
28 2
INTERNATIONAL JOURNAL
Group B-9 Dietz, Alma. Upjohn Co., Kalamazoo, Michigan] U . S. A. Wohicka, Wanda. P a h o w o w y Zaklad Higieny, U1. Chocimska 24, War szawa 12, Poland. Ohara, Y . and H. Nonomura. Faculty of Engineering] Yamanashi Univ., Kofu, Japan. GrOUD B- 10 Trejo. W. E. R. Squibb and Sons, New Brunswick, New J e r s e y , U. S. A. B r i n h a n n , Rolf. Organic Chem. Inst., Univ. of Gottingen, Gottingen, Germany Nishimura, H., M. Mayama and K. T a w a r a . Shionogi R e s . Lab., Shionogi and Co., Ltd., Fukushima-ku, Osaka, Japan.
Group B-11 Routien, J. B. and Corinne Clevenger. C h a r l e s P f i z e r and C o . , Inc. Groton, Connecticut, U. S. A. F a l c z o de M o r a i s , J.0, Inst. de Quimica d a Univ. Fed. d e Pernambuco, Recife, Pernambuco, Brazil. Okami, Y. Nat'1. Inst. of Health of Japan, Shinagawa, Tokyo, Japan. Group B- 1 2 Higgens, Calvin E. and R. E. Kastner. Eli Lilly and Co., Indianapolis, Indiana, U. S. A. W i l l i a m s , S. T. Hartley Botan. L a b o r . , Univ. of Liverpool, Liverpool, England. Vernon, T. R. Dept. Scientific and Ind. R e s . , P r i v a t e Bag, Auckland C1, New Zealand.
AGKNOW LEDGMENTS The Subcommittee on Actinomycetes of the Committee on Taxonomy, ASM, participated in t h e planning of t h e project, p r e p a r a t i o n of d e s c r i p tions and editing of t h e manuscript. M e m b e r s of the Subcommittee ( o t h e r than the authors) during the period covered by t h i s r e p o r t were: T.G. P r i d h a m , E.J. Backus, and W. Grundy. We a r e grateful t o A.A. Jchida, Jann M. Ichida, Janet Stevens, M a r g a r e t Speer, and Louise Hoffhines of the Ohio Wesleyan ISP staff f o r technical s e r v i c e s , f o r a s s i s t a n c e in locating authentic type c u l t u r e s and d e s c r i p t i v e l i t e r a t u r e , and for h e l p in collating data f r o m cooperating laboratories. C. W. C h r i s t e n s e n of Difco L a b o r a t o r i e s r e n d e r e d valuable a s s i s t a n c e in directing t h e formulation of dehydrated culture media according t o ISP specifications. The technical and financial a s s i s t a n c e of Difco Laborat o r i e s , Detroit,. Michigan, in supplying t h e s e standardized m e d i a t o coll a b o r a t o r s throughout the world is gratefully acknowledged.
SYSTEMATIC
BACTERIOLOGY
28 3
This r e s e a r c h was supported by National Science Foundation Grant GB 509 and GB 4365. The Subcommittee on Actinomycetes of the Committee on Taxonomy, ASM, and the Subcommittee on Taxonomy of Actinomycetes of the International Committee on Nomenclature of Bacteria a r e co- sponsoring advisors.
INT RODU C T ION This is the third in a s e r i e s of reports related to the comprehensive international effort to clarify taxonomy in the genera Streptomyces and Streptoverticillium through standardized collaborative study of the authentic type s t r a i n (or an acceptable neotype) for each of the named species. The cooperative study, known a s the International Streptomyces Project (ISP), is. sponsored by the Subcommittee onActinomycetes of the Committee on Taxonomy, ASM and the corresponding subcommittee of the International Committee on Bacteriological Nomenclature. The rationale, scope, extensive background preparation and general plan for the project a r e explained in detail in the f i r s t paper of the s e r i e s (Gottlieb and Shirling 1967); descriptive c r i t e r i a and methods for ISP characterization of Streptomyces and Streptoverticillium species a r e given under the title, Methods for Characterization of Streptomyces Species (Shirling and Gottlieb 1966). Illustrated descriptions for 100 type s t r a i n s o r suggested neotypes a r e given in the second paper of the s e r i e s (Shirling and Gottlieb 1968). This report ( P a r t I11 of the s e r i e s ) includes illustrated descriptions f o r an additional 100 type s t r a i n s o r suggested neotype strains f r o m the genus Streptomyces (including Actinomyces sensu Krasil'nikov) and the genus Streptoverticillium. The second paper of the s e r i e s (op. cit. pp. 73-79) should be consulted for analysis of a typical entry and guidance in interpretation of data submitted in each description. The ISP description for each species i s based upon data submitted by t h r e e independent collaborators. In r a r e instances where significant disagreement occurred in r e p o r t s f r o m the t h r e e laboratories, the cult u r e was either submitted to a fourth collaborator o r withdrawn for r e study. The ISP description i s an emendation to previous descriptions. The collaborators who studied the species a r e to be regarded a s the authors for the emendation. They a r e listed a s cooperating groups at the beginning of this paper and identified by group number in each description. E a r l i e r descriptions for each culture, including the description ascribed to the original author, a r e cited whenever possible to complete the characterization. The 200 type strains included in P a r t II and P a r t III of this s e r i e s a r e deposited a s reference cultures in the American Type Culture Collection, the Centraalbureau voor Schimmelcultures and the USSR Research Institute f o r Antibiotics. They will also be deposited in an appropriate culture collection in Japan. Cooperative studies characterizing approximately 300 o r m q r e additional species a r e in p r o g r e s s o r have been completed. Descriptions for these species will be included in subsequent r e p o r t s and the type s t r a i n s will be deposited in the culture collections a s the descriptions a r e published.
28 4
INTERNATIONAL
JOURNAL
ISP CHARACTERIZATIONS
OF TYPE STAINS'
Streptomyces aburaviensis Nishmura, Kimura, Tawara, Sa saki, Nakajima, Shimaoka, Okamoto, Shimohira and 1s.ono. Description: 1957, 205-211. Type strain: S-66 ISP 5033 Nishimuraet f r o m Nishimura a s S-66 = ISM 1 0 6 3 . ISP description by Group B - 1 . Spore chain morphology: Section Retiflexibiles (Fig. 1 ) . Mature s p o r e chains moderately long with LO to 50, o r s o m e t i m e s m o r e than 50, s p o r e s p e r chain. This morphology i s seen on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 2 ) . Color of colony: A e r i a l m a s s color in the Gray c o l o r - s e r i e s on y e a s t - m a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r a gine a g a r . R e v e r s e side of colony: Yellowish brown on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ; this pigment i s not a pH indicator. Color in medium: Melanoid pigments a r e not f o r m e d in peptoney e a s t - i r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment other than t r a c e of yellow i s found in medium in yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: Growth on carbon utilization media i s generally poor. _D-GLucose and _D-fructose and D_-xylose a r e utilized f o r growth. Reports varied f r o m no growth to t r a c e s of growth on L_-arabinose, s u c r o s e , 1-inositol, D_-mannitol, rhamnose and raffinose.
c.
w.).
S t r eptomy c e s aerocolonigene s Shinobu and Kawato. Description: ISP 5034 Shinobu and Kawato 1960a, 212-216. Type strain: 701 f r o m Shinobu a s 701. ISP description by Group B - 4 . Morphological c h a r a c t e r i s t i c s : No a e r i a l mycelium i s f o r m e d on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . In addition to these media, c o o p e r a t o r s found no a e r i a l mycelium o r s p o r e formation on Langeronls weak potato-carrot medium, modified Bennett's medium, tomato-paste oatmeal a g a r , soil e x t r a c t a g a r , modified Sabouraud maltose a g a r o r Czapek's a g a r plus glucose. Spore s u r face: Not observed. The original description by Shinobu (tbld.) includes the following statement: "No. 701 had a tendency to become weak and .I1. Lose i t s formation ability of a e r i a l mycelia by successive cultures,. Shinobu observed b e s t a e r i a l mycelium development on glucose-asparagine a g a r . On this medium he observed flexuous, cottony white a e r i a l mycelium without whirls o r s p i r a l s and with spherical to oval conidia. Special morphological c h a r a c t e r i s t i c s : Although ISP collaborators w e r e unable to obtain a e r i a l mycelium on any medium t r i e d (see above), Shinobu's original description makes special mention of t h e formation of secondary colonies on the cottony a e r i a l mycelium on glucose-asparagine a g a r . His illustrations suggest that these seconday colonies a r e s i m i l a r to llsclerotia'l r e p o r t e d by other a u t h o r s (Waksman 1961, 292).
w.).
..
I Type s t r a i n s for LOO Streptomyces s p e c i e s a r e described h e r e . An additional LOO s t r a i n s w e r e d e s c r i b e d i n an e a r l i e r r e p o r t (og. c i t . ) ; and descriptions f o r other species a r e in preparation.
F i g u r e 1. 2. aburaviensis. R F s p o r e chains. (a) ( X 100) on oatmeal agar, 24 days. (b) (X 640)on s a l t s - s t a r c h agar, 14 days. F i g u r e 2. 2. aburaviensis. Smooth spores; electron micrograph from 14 day culture on oatmeal a g a r . Figure 3 . S. afghaniensis. Spiral spore chains (X 400). Short chains f r o m incomplete s p i r a l s or hooks. afghaniensis. Spiny spores; electron micrograph from F i g u r e 4. 14 day culture on oatmeal a g a r . *
5.
Sources of illustrations a r e indicated by s u p e r s c r i p t number following legend for each figure, and a r e listed by number after the bibliography.
28 6
INTERNATIONAL
JOURNAL
R e v e r s e side of colony: No distinctive pigments (pale grayed yellow)

on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r g l y c e r o l - a s p a r -
agine a g a r . Color i n medium: Melanoid pigments a r e not f o r m e d on peptoney e a s t - i r o n a g a r , tyrosine a g a r o r peptone-yeast broth. P i g m e n t s other than t r a c e s of yellow a r e not found in the medium in y e a s t - m a l t a g a r , oatmeat a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: D_-Glucose, L_-arabinose, s u c r o s e , Q-xylose, i inositol, -Dmannitol, p f r u c t o s e and rhamnose a r e utilized f o r growth. Growth on rhamnose is Less abundant than on the other carbon s o u r c e s . No growth o r only t r a c e of growth on raffinose. Streptomyces afghaniensis Shimo, Shiga, Tomosugi and Kamoi. Description: Shimo e t a1 . 1959, 1-6. Type strain: 772 (Kd.). ISP 5528 f r o m Shirno as 772. 1SP description by Group B-5. Spore chain morphology: Section S p i r a l e s , but s h o r t s p o r e chains usually r e s u l t in hooks, incomplete s p i r a l s o r s p i r a l s with only one o r two t u r n s (Fig. 3 ) . Short, s t r a i g h t and flexuous chains of only a few s p o r e s a r e common, but hooks a n d loops of wide d i a m e t e r a s found in typical c u l t u r e s a r e not found. Spore chains a r e short, often with only 3 to 10 s p o r e s p e r chain, but m o r e than 10 s p o r e s may be found on some chains. This morphology i s b e s t developed on oatmeal a g a r a n d s a l t s - s t a r c h a g a r . Spore surface: Spiny (Fig. 4). Color of colony: A e r i a l m a s s color in the Blue c o l o r - s e r i e s on oatm e a l a g a r . I m m a t u r e aerial mycelium o r mycelium producing few s p o r e s is in the Yellow c o l o r - s e r i e s . O b s e r v e r s r e p o r t e d both Blue and Yellow c o l o r - s e r i e s f o r y e a s t - m a l t a g a r a n d s a l t s - s t a r c h a g a r . Mature s p o r e s a r e not usually found on glycerol-asparagine a g a r . R e v e r s e side of colony: Grayed yellow on oatmeal a g a r , modified to orange o r reddish brown on y e a s t - m a l t a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ; this pigment i s not a pH indicator. Color in medium: Melanoid pigments a r e f o r m e d in peptone-yeasti r o n a g a r , tyrosine a g a r and peptone-yeast broth. Orange to r e d o r brown pigments a r e found i n the medium in y e a s t - m a l t agar, oatmeal a g a r , s a t t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . These pigments a r e not pH sensitive when tested with 0.05 N H C l o r NaOH. Carbon utilization: E G l u c o s e , L_-arabinose, sucrose, D_-xylose, i -inositot, D_-mannitol, Q-fructose, rhamnose and raffinose a r e utilized f o r growth. Streptomyces atbidofuscue Maeda, Kosaka, Okami and Umezawa 1953. Streptomyces albidofuscus Okami and Umezama Okami, Maeda and Umezawa 1954. The type s t r a i n is d e s c r i b e d under the n a m e Streptomyces pyridomyceticus Okami and Umezawa, 9:~. Actinomyces albocyaneus Krasil'nikov and Agre. Description: Krasil'nikov and Agre 1960, 272. Type s t r a i n : INMI 679 (2). ISP 5197 f r o m Krasil'nikov a s INMI 679 = RI A - 6 3 3 . ISP description by Group B -9. Spore chain morphology: Section Retinaculiaperti o r Spira les. Spirals, when formed, a r e open a n d poorly developed. Long chains often show primitive s p i r a l s , t e r m i n a l s p i r a l s , loops o r hooks c h a r a c t e r i s t i c of % c u l t u r e s (Fig. 5). Flexuous chains a r e a l s o common.
F i g u r e 5. A. albocyaneus. R A or Spiral spore chains (X 1100).5 (a) FlTmous looped a n d p i r a l e d i n s on yeast-malt a g a r , 14 days. (b) Open s p i r a l s , yeast-malt agar, 14 days. (c) Flexuous chains and t e r m i n a l spirals, oatmeal a g a r , 21 days. F i g u r e 6. _A. albocyaneus. Spiny spores, s h o r t spines; electron micrograph f r o m 10 day culture on yeast-malt a g a r . F i g u r e 7. 2. albofaciens. Spiral s p o r e chains (X 1100). (a) On oatmeal agar, 21 days. (b) On glycerol-asparagine a g a r , 1 4 days. F i g u r e 8. 2. albofaciens. Smooth s p o r e s ; electron micrograph f r o m 14 day culture on glycerol-asparagine a g a r .
'
28 8
INTERNATIONAL
JOURNAL
Mature s p o r e chains generally m o r e than LO but l e s s than 50 s p o r e s p e r chain on y e a s t - m a l t a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r . Spore surface: Spiny; spines a r e usually s h o r t and thick (Fig. 6). Special morphological c h a r a c t e r i s t i c s : Fragmentation of the subs t r a t e mycelium on s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r to f o r m I1oidiosporesl1is r e p o r t e d by one o b s e r v e r ; a second o b s e r v e r r e c o r d s 'lchlamydosporesll on oatmeal a g a r and glycerol-asparagine a g a r a f t e r 7 days. Color of colony: A e r i a l m a s s color in the G r e e n c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r ; a e r i a l myceltium i s poorly developed o r white on glycerol-asparagine a g a r . R e v e r s e side of colony: Grayed yellow is modified by g r e e n o r blue pigments on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Only one o b s e r v e r r e c o r d e d a change i n r e v e r s e color f r o m pale g r e e n to pale violet by addition of 0.05 N HC1. Color in medium: Melanoid pigments a r e not found i n peptone-yeast iron a g a r , tyrosine a g a r o r tryptone-yeast broth. N o pigment i s found in the medium in y e a s t - m a t t a g a r , s a l t s - s t a r c h a g a r o r g l y c e r o l - a s p a r a gine a g a r . Carbon uti t i za tion : R-G luco s e , &-a rabinos e , s u c r o s e , _D -xy 10se , i -inositol, _D -mannitol, _D-fructrose, rhamnose a n d raffinose a r e utilized for growth, although r e p o r t s v a r y f r o m slight growth to good growth f o r s u c r o s e , &-inositol and raffinose. Streptomyces albofaciens T h i r u m a l a c h a r and Bhatt. Description: ISP Thirumalachar and Bhatt 1960, 61-63. Type strain: 27-A 5268 f r o m M. J . Thirumalachar a s 27-A. ISP description by Group B-9. Spore chain morphology: Section Spirales (Fig. 7). Open i r r e g u l a r s p i r a l s s o m e t i m e s a p p e a r to a r i s e f r o m a n axial hypha, but t r u e whorls typical of verticillate c u l t u r e s a r e not f o r m e d . Mature s p o r e chains generally LO to 50, o r s o m e t i m e s m o r e than 5 4 s p o r e s p e r chain. This morphology is s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 8). Color of colony: A e r i a l m a s s color in the White c o l o r - s e r i e s on y e a s t - m a l t a g a r and glycerol-asparagine a g a r ; White o r G r a y color - s e r i e s on oatmeal a g a r and inorganic s a l t s - s t a r c h a g a r . R e v e r s e side of colony: N o distinctive pigments on yeast-malt a g a r , oatmeat a g a r , s a l t s - s t a r c h a g a r o r glycero 1-asparagine a g a r . Color i n medium: Melanoid pigments a r e not f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment o r only t r a c e of yellow pigment, found in medium i n yeast-matt a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r o r glycero 1-asparagine a g a r . Carbon uti Lization: &Glucose, 4 - a r a b i n o s e , - i n o s i t o l , _D-mannitol, I l f r u c t o s e and raffinose a r e utilized f o r growth. No growth o r only t r a c e of growth on rhamnose. Utilization of s u c r o s e and _D-xylose i s doubtful.
w.).
Streptomyces alboflavus (Waksman and C u r t i s ) Waksman and Henrici. Descriptions: Actinomyces albo-flavus Waksman and C u r t i s 1916, 120 and 128; Actinomyces atbofkivue Waksman 1919, 90-91; Streptomyces alboflavus (Waksman a n d C u r t i s ) Waksman and Henrici 1948, 954. Type strain: IMRU 3008 (Waksman 1961, 169 and 170). ISP 5045 f r o m S. A. Waksman a s IMRU 3008. ISP description by Group B-5.
SYSTEMATIC
BACTERIOLOGY
28 9
Morphological c h a r a c t e r i s t i c s : Typical a e r i a l mycelium i s not f o r m e d on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerola s p a r a g i n e a g a r . Spore chain morphology, s p o r e surface and a e r i a l m a s s color of colony cannot be observed on ISP media. L o s s of ability to produce sporulating a e r i a l mycelium was noted in a n e a r l y description of this culture (Waksman 1919, 90). E a r l y descriptions of the culture d e s c r i b e white o r yellowish white a e r i a l mycelium on synthetic a g a r o r Czapekls a g a r only. Special morphological c h a r a c t e r i s t i c s : Coremia formation on s a l t s s t a r c h a g a r and glycerol-asparagine a g a r was recorded by two o b s e r v e r s . This s a m e phenomenon was recorded in the original description on a e r i a l mycelium was found Czapekls a g a r (Waksman 1916, 120) : ' I . a m a s s of hyphae m a s s e d to have a tendency to p r o d u c e , together into a rope, and f r o m this rope fine filaments coming out i n the shape of side branches. The s t r u c t u r e looks like the root of a t r e e and fine rootlets coming out on the side." One ISP o b s e r v e r found straight s p o r e chains within the coremia on glycerol-asparagine a g a r . R e v e r s e side of colony: N o distinctive pigment (grayed yellow o r grayed greenish yellow) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Color i n medium: Melanoid pigments f o r m e d weakly in peptone y e a s t - i r o n a g a r and tryptone-yeast broth but not i n tyrosine a g a r . P i g ments other than melanoids not f o r m e d in yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D -G luco s e , ,L -arabinose, g x y l o se, i - i n 0 sitol, D -mannitol, _D -fructose and raffinose a r e utilized f o r growth. No growth o r only t r a c e of growth on rhamnose. Utilization of s u c r o s e i s doubtful.
. .. . .
..
Streptomyces albogriseolus Benedict, Shotwell, Pridham, Lindenf e l s e r and Haynes. Description: Benedict e t a l . 1954, 653-656. Type s t r a i n : 7-A = NRRL B -1305 k d . ) . ISP 5003 f r o m T. P r i d h a m a s 7-A = NRRL B-1305. Note: The authenticity of this s t r a i n a s the type s t r a i n deposited by Dr. R . G . Benedict in the ARS Culture Collection in August, 1951, a s NRRLB-1305 has been verified by C. W. Hesseltine and T. P r i d h a m of the ARS Culture Collection, P e o r i a , 111. (personal communication). No authentic subcultures f r o m the type s t r a i n have hairy s p o r e s . At some t i m e the culture with hairy s p o r e s was apparently mislabelled a s NRRL B-1305 and given limited distribution. The warty to smooth spored ISP 5003 (7-A = NRRL B-1305) c o r r e s p o n d s to the original d e s cription i n a l t details. The hairy spored culture was a l s o studied by ISP collaborators a s ISP 5344. In addition to the difference i n s p o r e surface, i t differs f r o m the type s t r a i n and the original description i n carbon utilization. The type strain, i n a g r e e m e n t with the description by Benedict e t a t . , u s e s a l l ISP carbon s o u r c e s except raffinose and cellulose. The h a i r y spored s t r a i n i s unable to utilize any of the ISP carbon s o u r c e s except _D - glucose and possibly fructose. ISP d e s c r i p tion by Group A-2. Spore chain morphology: Section Spirales. S p i r a l s a r e open (Fig. $9); s p o r e chains a r e a l s o common. Mature spore chains flexuous o r generally LO to 50 s p o r e s p e r chain. This morphology i s seen on yeastm a l t a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r , but not on glycerola s p a r a g i n e a g a r . Spore surface: Warty. Warts a r e not prominent o r r e g u l a r and some smooth s p o r e s may be fouud (Fig. LO).
29 0
Color of colony: A e r i a l m a s s color in the Gray c o l o r - s e r i e s on yeast-matt a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r . R e v e r s e side of colony: No distinctive pigments a r e f o r m e d (colorl e s s o r yellowish gray on oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerola s p a r a g i n e a g a r ; grayed yellowish brown o r olive brown on y e a s t - m a l t agar). Color in medium: Melanoid pigments a r e not f o r m e d i n peptoney e a s t - i r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment i s found in medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D -Glucose, s u c r o s e , _D xylose, i - i n o s i t o l , Dmannitol, P f r u c t o s e and rhamnose a r e utilized f o r growth, but growth on s u c r o s e is l e s s abundant than on other carbon s o u r c e s . No growth o r only t r a c e of growth on raffinose. Streptomyces a m a k u s a e n s i s Nagatsu, Ansai, Ohkuma and Suzuki. Description: Nagatsu et a t . 1963, 207-210. Type s t r a i n : 10-101 ( i x d .). ISP 5219 f r o m S . Suzuki, Inst. Phys. and Chem. R e s . , Tokyo a s LO-101. ISP description by Group B-3. Spore chain morphology: Section S p i r a l e s (Fig. 11). Mature s p o r e chains 10 to 50 o r m o r e s p o r e s p e r chain on y e a s t - m a l t a g a r a n d s a l t s s t a r c h a g a r . Sporulating a e r i a l mycelium i s poorly developed o r a b s e n t on oatmea 1 a g a r and glycerol-asparagine a g a r . Spore surface: Srnooth (Fig. 12). Color of colony: Aerial m a s s color of m a t u r e a e r i a l mycelium i s in the Blue c o l o r - s e r i e s on s a l t s - s t a r c h a g a r and the G r e e n o r Blue c o l o r - s e r i e s on yeast-matt a g a r ; younger mycelium may be in the G r a y c o l o r - s e r i e s . Mature a e r i a l mycelium i s usually not f o r m e d on oatmeal a g a r o r glycerol-asparagine a g a r . R e v e r s e side of colony: No distinctive pigments (colorless o r pale yellow) on y e a s t - m a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerolasparagine agar. Color i n medium: Melanoid pigments a r e usually f o r m e d i n peptoneyeast-iron a g a r , but not in tyrosine a g a r . N o pigments a r e found i n the medium in yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerolasparagine agar. Carbon utiliwtion: This culture does not show good growth with any of the carbon s o u r c e s t e s t e d on P r i d h a m and Gottlieb carbon utilization medium. _D -Glucose i s utilized f o r growth. No significant growth o c c u r s on _D -xylose, i -inositol, _D -mannitoL, P f r u c t o s e , r h a m n o s e o r raffinose and only doubtful t r a c e i s s e e n on ,L -arabinose and s u c r o s e . Streptomyces ambofaciens Pinnert-Sindico. Description: P i n n e r t Sindico 1954, 702-707. Type s t r a i n : 3486 ISP 5053 f r o m R. Despois, Soci6ZC Rh6ne-Poulenc a s 3486. ISP description by Group B -8. Spore chain morphology: Section S p i r a l e s (Fig. 13). Open t e r m i n a l s p i r a l s on long s p o r e chains a l s o s u g g e s t E morphology. Mature s p o r e chains generally LO t o 50 o r m o r e s p o r e s p e r chain. This m o r phology i s seen on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r , although sporulating a e r i a l mycelium i s not abundant on glycerol-asparagine a g a r . Spore surface: Smooth to w a r t y ; surface i r r e g u l a r i t i e s suggesting w a r t s a r e s m a l l (Fig. 14).
m.).
F i g u r e 9 . 2. albogriseolus. ( a , b) S p i r a l s p o r e chains ( X 500) on g l y c e r o l - a s p a r a g i n e a g a r , 7 days.8 F i g u r e 10. S. albogriseolus. Warty and smooth s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 21 day c u l t u r e on y e a s t - m a l t a g a r . F i g u r e 11. S. a m a k u s a e n s i s . ( a ) Open and tight s p i r a l s (X 300) on y e a s t m a l t a g a r , 14 days. (b) Tight s p i r a l s , y e a s t - m a l t a g a r , 15 days. F i g u r e 12. S . a m a k u s a e n s i s . Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 14 day c u l t u r e on y e a s t - m a l t a g a r . l 2 F i g u r e 13. S. ambofaciens. S p i r a l s p o r e chains ( X 800) on s a l t s - s t a r c h agar, 1 4 d a y s . F i g u r e 14. S. ambofaciens. Smooth s p o r e s with m i n o r s u r f a c e i r r e g u laritiessuggesting small warts o r spines; electron micrograph from 14 day c u l t u r e on y e a s t - m a l t a g a r . I 4
'"
''
29 2
INTERNATIONAL
JOURNAL
Color of colony: A e r i a l m a s s c o l o r i n the G r a y c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r and salts-starch a g a r . R e v e r s e s i d e of colony: No distinctive pigments ( c o l o r l e s s to g r a y e d yellow) on y e a s t - m a l t a g a r , o a t m e a l a g a r and g l y c e r o l - a s p a r a g i n e a g a r . Dark brown, d a r k blue or a l m o s t black s u b s t r a t e m y c e l i u m pigment i s found on s a l t s - s t a r c h a g a r . This pigment i s not pH s e n s i t i v e . Color in m e d i u m : Melanoid pigments a r e not f o r m e d i n peptoney e a s t - i r o n a g a r , t y r o s i n e a g a r or tryptone-yeast broth. Yellow pigment m a y be found i n m e d i u m i n s a l t s - s t a r c h a g a r ; no pigment i s i n the m e d i u m i n y e a s t - m a l t a g a r , o a t m e a l a g a r or g l y c e r o l - a s p a r a g i n e a g a r . Carbon utilization: _D -Glucose, 4 - a r a b i n o s e , _D -xylose, i -inositol, D -mannitol, _D - f r u c t o s e and r h a m n o s e a r e utilized f o r growth. NO growth or only t r a c e of growth on raffinose. Utilization of s u c r o s e i s doubtful.
S t r e p t o m y c e s aminophilus Wooldridge. Description: The n a m e a n d oriRinal identification f o r t h i s s p e c i e s i s c r e d i t e d to J . W. F o s t e r by Oswald e t a t . 1956, 236. This publication contains no d e s c r i p t i o n and the o r i g i n a l c h a r a c t e r i z a t i o n s by F o s t e r have been lost (J. W. F o s t e r , p e r s o n a l communication). The f i r s t available d e s c r i p t i o n i s i n Woold r i d g e 1957, G e r m a n P a t e n t Notation DAS 1000966; s e e a l s o Wooldridge 1960a, B r i t i s h P a t e n t 828, 792 and Wooldridge L960b, U. S . P a t e n t 2,956,925 ( S t r e p t o m y c e s c o e l i c o t o r v a r . aminophilus). Type s t r a i n : NRRL 2390 (Ibld.; L. E. Arnow, W a r n e r - L a d e r t R e s . I n s t . , p e r s o n a l communication). ISP 5186 f r o m C. W. H e s s e l t i n e a s N R R L 2390. ISP d e s c r i p t i o n by G r o u p A-13. S p o r e chain morphology: Section S p i r a l e s (Fig. 15). Mature s p o r e chains 3 t o 10 or s o m e t i m e s m o r e than 10 s p o r e s p e r chain. T h i s m o r phology is s e e n on y e a s t - m a l t a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r a g i n e a g a r , but f o r m a t i o n of sporulating a e r i a l m y c e l i u m i s poor on o a t m e a l a g a r . Spore s u r f a c e : Smooth ( F i g . 16). Color of colony: A e r i a l m a s s c o l o r i n the Yellow c o l o r - s e r i e s on yeast-malt a g a r , salts-starch a g a r and glycerol-asparagine a g a r ; a e r i a l m y c e l i u m poorly developed on o a t m e a l a g a r . R e v e r s e s i d e of colony: No distinctive pigments (yellow or g r a y e d yellow) on y e a s t - m a t t a g a r , o a t m e a t a g a r , s a l t s - s t a r c h a g a r or g l y c e r o l asparagine agar. Color i n medium: Melanoid pigments not f o r m e d i n peptone-yeasti r o n a g a r , t y r o s i n e a g a r o r t r y p t o n e - y e a s t broth. N o pigments, o r only t r a c e s of yellow pigment, a r e found i n the m e d i u m i n y e a s t - m a l t a g a r , oatmeal a g a r , salts-starch a g a r o r glycerol-asparagine a g a r , Carbon utilization: _D -Glucose, _D -xylose, _D -mannitol a n d _D f r u c t o s e a r e utilized f o r growth. No growth o r only t r a c e of growth on i -inositol and r h a m n o s e . R e p o r t s v a r y on utilization of s u c r o s e a n d raffinose.
(Waksman and Woodruff) Waksman a n d H e n r i c i . Description: Actinomyces antibioticus Waksman and Woodruff 1941, 246-249; S t r e p t o m y c e s antibioticus (Waksman and Woodruff) Waksman a n d H e n r i c i 1948, 942. Type s t r a i n : IMRU 3435 (Waksman 1961; R. Gordon, P e r s o n a l Communication). I S P 5234 f r o m S.A. Waksman a s IMRU 3435. ISP d e s c r i p t i o n by Group B-10.
Figure 15. 2. aminophilus. Spiral s p o r e chains ( X 500) on s a l t s - s t a r c h agar, 23 days.15 Figure 16. S. aminophilus. Smooth s p o r e s ; electron micrograph from 1 5 day culture on yeast-malt a g a r . S. antibioticus. R F s p o r e chains (X 600). l 6 F i g u r e 17. F i g u r e 18. 2. antibioticus. Smootb s p o r e s ; electron micrograph f r o m 21 day culture on yeast-malt a g a r . F i g u r e 19. S. antimycoticus. Compact c l u s t e r s of closed s p i r a l s on s a l t s - s t a r c h agar, 14 days ( X 200)." F i g u r e 20. S. antimycoticus. Spiny t o warty s p o r e s in s p i r a l s p o r e chains; e l e c t r o n micrograph f r o m 14 day culture on s a l t s - s t a r c h agar. "
''
29 4
INTERNATIONAL
JOURNAL
Spore chain morphology: Section Rectiflexibiles (Fig. 17). Mature s p o r e chains a r e short, often with only 3 to 10 s p o r e s p e r chain, longer chains a r e a l s o found. This morphology is observed on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 18). Color of colony: A e r i a l m a s s color in the G r a y c o l o r - s e r i e s on yeast-malt a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ; m a t u r e a e r i a l mycelium poorly developed on o a t m e a l a g a r . R e v e r s e side of colony: N o distinctive pigments (grayed yellowbrown on y e a s t - m a l t a g a r , grayed yellow o r d a r k g r e y i s h greenish yellow on oatmeal a g a r , s a l t s - s t a r c h a g a r , glycerol-asparagine a g a r ) ; s u b s t r a t e mycelium pigment is not a pH indicator. Color i n medium: Melanoid pigments a r e produced in peptone-yeasti r o n a g a r , tyrosine a g a r and peptone-yeast broth. No pigment i n medium, f yellow, i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h o r only t r a c e o a g a r o r glycerol-asparagine a g a r . Carbon utilization: ,D -Glucose, &-arabinose, i - i n o s i t o l , _D-mannitol, D-fructose and rhamnose a r e utilized f o r growth. No growth o r only t r a c e of growth on s u c r o s e and raffinose. Utilization of _D-xylose i s doubtful. Streptomyces antimycoticus Waksman, Descriptions:
ISP 5284 f r o m C. W. Waksman 1957, 799. Type s t r a i n : A158 Hesseltine a s N R R L 2421 = A158. ISP description by Group B-11. Spore chain morphology: Section Spirales; compact c l u s t e r s of closed s p i r a l s on yeast-malt a g a r , oatmeal a g a r , salts-starch a g a r a n d glycerol-asparagine a g a r (Fig. 19); m a t u r e s p o r e chains of generally LO to 50 s p o r e s p e r chain on these media. Spore surface: Spiny to warty. Surface i r r e g u l a r i t i e s on s p o r e s a r e intermediate between v e r y short, thick spines and w a r t s (Fig. 20). Color of colony: A e r i a l m a s s color i n the G r a y c o l o r - s e r i e s on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . R e v e r s e side of colony: No distinct pigment (pale grayed yellow o r grayed greenish yellow) on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e not produced on peptoneyeast-iron a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment i s found in medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-Glucose, &-arabinose, s u c r o s e , _D-xylose, i -inositol, _D-mannitol, _D-fructose, rhamnose and raffinose a r e utilized for growth. Growth on _L-arabinose and 2 - x y l o s e i s generally l e s s abundant than OD the other carbon s o u r c e s .
Streptomyces a r a b i c u s Shibita, Nakazawa, Miyake, Inoue and Okabori. Description: Shibita et a l . 1957, 32-37. Type s t r a i n : ISP 5252 f r o m K. Nakazawa a s I F 0 3406 = 6762. ISP 6762 description by Group B-8. Spore chain morphology: Section S p i r a l e s on oatmeal a g a r a n d s a l t s - s t a r c h a g a r (Fig. 2La). Short s p o r e chains on yeast-malt a g a r and glycerol-asparagine a g a r may appear to be RF morphology o r may
9 Leben, S t e s s e l and Keitt 1952, 159-168; Streptomyces antimycoticus
Streptomyce s
m).
m.).
SYSTEMATIC
BACTERIOLOGY
29 5
show only incomplete s p i r a l s or hooks (Fig. 21b). Mature s p o r e chains on oatmeal a g a r and s a l t s - s t a r c h a g a r generally LO to 50 or m o r e s p o r e s p e r chain; s h o r t e r s p o r e chains a r e found on yeast-malt a g a r and glycerol-asparagine a g a r . Spore surface: Warty to spiny. Surface i r r e g u l a r i t i e s a r e s h o r t and blunt (Fig. 22a); smooth s p o r e s may a l s o be found (Fig. 22b). Color of colony: A e r i a l m a s s color i n the G r a y c o l o r - s e r i e s on y e a s t - m a l t a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r ; Red c o l o r - s e r i e s on glycerol-asparagine a g a r . O b s e r v e r s selected color tabs r e p r e s e n t ing neutral g r a y s to light grayish reddish brown (5fe) for the Gray colors e r i e s and grayish yellowish pink (5dc) f r o m the Red c o l o r - s e r i e s for glycerol-asparagine a g a r . R e v e r s e side of colony: G r a y i s h yellowish brown on yeast-malt a g a r ; c o l o r l e s s to pale yellowish g r a y o r brown on a a t m e a l a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Substrate mycelium pigment i s not a pH indicator. Color in medium: Melanoid pigment is not produced in peptone-yeasti r o n agar, t y r o s i n e a g a r , o r tryptone-yeast broth. T r a c e of yellow pigment m a y be found in the medium i n o a t m e a l a g a r or s a l t s - s t a r c h a g a r or m a y be absent. This pigment is not pH sensitive. Carbon utilization: D- Glucose, 2 - a r a b i n o s e , D-xylose, i=inositol, D-mannitol, _D-fructose and rhamnose a r e utilized f o r growth. No growth or only trace of growth on raffinose. Utilization of s u c r o s e is doubtful. Streptomyces argenteolus F r i e d , P e r l m a n , Langlykke and Titus. Descriptions: Actinomycete s p e c i e s ATCC 11,009 P e r l m a n e t at. US. P a t e n t 2, 709, 705, 1955; Streptomyces argenteolus ATCC 11,009, F r i e d e t a t . U.S. Patent 2,855, 343 Oct., 1958; Streptomyces argenteolus (name only) T r e s n e r e t at. 1961, 74. Type strain: ATCC 11,009 k d . ) . ISP 5226 f r o m F. Arnow, Squibb Culture Collection a s MD 2428 = ATCC 11,009. ISP description by Group B-3. Spore chain morphology: Section Spirales (Fig. 23). This culture is described a s Itnot forming loops o r spirals" i n the original d e s c r i p tions appearing i n the patents cited above. However, the type s t r a i n was observed to produce s p i r a l s i n subsequent studies by P r i d h a m e t at., 1958 and by T r e s n e r et at., 1961. Mature s p o r e chains generally 10 t o 50 o r m o r e s p o r e s p e r chain on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 24). Color of colony: A e r i a l m a s s color i n the G r a y c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . R e v e r s e side of colony: Colorless to grayish yellowish g r e e n o r g r a y on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerola s p a r a g i n e a g a r ; s u b s t r a t e mycelium pigment i s not a pH indicator. Color i n medium: Melanoid pigments not f o r m e d in peptone-yeasti r o n a g a r , tyrosine a g a r or tryptone-yeast broth. No pigment found i n medium i n yeast-malt a g a r , oatmeal a g a r , s b l t s - s t a r c h a g a r or g Lycerol-asparagine a g a r . Carbon utilization: _D-Glucose, &-arabinose, _D-xylose. _D-mannitol, D-fructose a n d rhamnose a r e utilized f o r growth. N o growth or only t r a c e of growth on sucrose, i - i n o s i t o l and raffinose.
--
--
F i g u r e 21. S. a r a b i c u s . ( a ) S p i r a l s p o r e c h a i n s ( X 300) o n s a l t s - s t a r c h agar, ( b ) Short, flexuous chains with s o m e hooks and incomplete s p i r a l s ( X 800) on y e a s t - m a l t a g a r , 14 days. l 3 F i g u r e 2 2 . S. a r a b i c u s . (a) Spiny t o w a r t y s p o r e s ; s u r f a c e i r r e g u l a r i t i e s a r e thick a n d o r rounded.19 ( b ) Smooth s p o r e s of S. a r a b i c u s ; e l e c t r o n m i c r o g r a p h f r o m 1 4 day c u l t u r e on y e a s t - m a l t a g a r . I 4 F i g u r e 2 3 . 2. a r enteolus. S p i r a l s p o r e chains ( X 600) on y e a s t - m a l t a g a r , 14 days.
--%---
F i g u r e 24. S. a r g e n t e o l u s . Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 14 day c u l t u r e on o a t m e a l a g a r . lo
SYSTEMATIC
BACTERIOLOGY
29 7
Actinomyces aurantiogriseus Preobrazhenskaya. Description: P r e o b r a z h e n s k a y a k Gauze e t a t . 1957, 74. Type strain: 10369/58 (G.F. Gauze, cited by Gottlieb 1968, 20). Streptomyces aurantiogriseus (Preobrazhenskaya) P r i d h a m e t at. 1958, 67. ISP 5138 f r o m T.P. Preobrazhenskaya a s 10369/58. ISP description by Group B - 6 . Spore chain morphology: Section Retinaculiaperti o r Spirales. S p i r a l s a r e open and often a r e i r r e g u l a r and poorly developed (Fig. 25a, b). Long s p o r e chains of the = t y p e a n d flexuous chains a r e common (Fig. 25c). Mature s p o r e chains may be slow to develop; they a r e moderately long with m o r e than.10 s p o r e s p e r chain on oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 26). Color of colony: A e r i a l m a s s color i s i n both the Red and the G r a y c o l o r - s e r i e s on yeast-malt a g a r , s a l t s - s t a r c h a g a r and glycerolasparagine a g a r ; both c o l o r s may a p p e a r on the s a m e medium. One o b s e r v e r noted an i n c r e a s e i n g r a y a e r i a l color between 14 a n d 21 days on glycerol-asparagine a g a r ; this tendency to change f r o m r e d to g r a y was a l s o included i n the original description. The color tabs m o s t frequently chosen w e r e 2ec (yellowish gray) f r o m the G r a y c o l o r - s e r i e s and 5ge (tight grayish reddish brown) f r o m the Red c o l o r - s e r i e s . R e v e r s e side of colony: No distinctive pigment (grayed yellow on oatmeal a g a r and glycerol-asparagine a g a r to orange-yellow o r brown on yeast-malt a g a r and s a l t s - s t a r c h agar). Substrate mycelium pigment is not a pH indicator. Color i n medium: Melanoid pigments a r e produced i n peptone-yeasti r o n a g a r , tyrosine a g a r and tryptone-yeast broth. Pigments other than melanoids a r e not found i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine agar. Carbon utilization: _D-Glucose, &-arabinose, sucrose, _D-xylose, i -inositol, _D-mannitol, _D-fructose, rhamnose and raffinose a r e utilized f o r growth. St r ept omyce s a u r e ofa ciens Dugga r De sc ription: Dug ga r 1948, 177 181; U. S . P a t e n t 2,482,055 Sept. 1949. Type strain: A-377 = NRRL 2209 k d . ) . ISP 5127 f r o m E. Backus, Lederle Labs., A m e r . Cyanamid Co. a s A-377 = NRRL 2209. ISP description by Group A-12. Spore chain morphology: Section Retinaculiaperti but chains r e p r e sentative of Section K a r e a l s o common (Fig. 27). Mature s p o r e chains generally LO to 50 o r m o r e s p o r e s p e r chain. This morphology i s seen on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerola s p a r a g i n e a g a r . Spore surface: Smooth (Fig. 28). Color of colony: A e r i a l mass color i n the G r a y c o l o r - s e r i e s on yeastm a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . R e v e r s e side of co tony: No distinctive pigments (grayed yellow, orange-yellow o r brown on yeast-malt a g a r , and grayed yellow o r greeni s h yellow on oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ) . Substrate myce Lium pigment i s not a pH indicator. Color i n medium: Melanoid pigments not f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. N o pigment i s found i n medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: D_-Glucose, &-arabinose, sucrose, _D-xylose and D-fructose a r e utilized f o r growth. No growth o r only t r a c e of growth on&-inositol, _D-mannitol, rhamnose and raffinose.
F i g u r e 25. A. a u r a n t i o g r i s e u s . (.a) Incomplete s p i r a l s o r hooks ( X 500) on y e a s t - m a l t a g a r , 2 3 d a y s . l5 (b) S p i r a l s ( X 700) on y e a s t - m a l t a g a r , 2 1 d a y s . " (c) Flexuous s p o r e c h a i n s ( X 350) o n y e a s t - m a l t agar, 18 d a y s . " F i g u r e 26. A. a u r a n t i o g r i s e u s . Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 1 8 2 a y culture on yeast-malt a g a r . " F i g u r e 27. S. a u r e o f a c i e n s . RA s p o r e c h a i n s ( X 240) on o a t m e a l a g a r , 14 days? F i g u r e 28. S. a u r e o f a c i e n s . Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h from 14 day c u l t u r e on o a t m e a l agar. F i g u r e 29. b a a r n e n s i s . Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h from 14 day c u l t u r e on g l y c e r o l - a s p a r a g i n e a g a r .
2.
''
'
SYSTEMATIC
BACTERIOLOGY
299
(Duchg) Pridham, Hesseltine and Benedict. Descriptions: Actinomyces v i r i d i s Duchd 1934, 311-317; Actinomyces b a a r n e n s i s Duchd 1934 = Actinomyces v i r i d i s Gougerot, B lum and Duchk 1934, 376. Type strain: Dreyfus 472 (G.A. deVries, CBS, .p e r s o n a l communication). ISP 5232 f r o m G.A. d e V r i e s a s Dreyfus 472. ISP description by Group B-1. Spore chain morphology: This s t r a i n h a s apparently lost the ability to produce good sporulating a e r i a l mycelium. Spore chains, when found, w e r e straight (Section Rectiflexibiles) and usually contained m o r e than LO s p o r e s p e r chain. Spore surface: Smooth (Fig. 29). Color of colony: A e r i a l mycelium on yeast-matt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r i s inadequate f o r d e t e r mination of a e r i a l m a s s color. The original description of Duchd h.2.) d e s c r i b e s e a r l y appearance of abundant greenish a e r i a l myce lium on comparable media. R e v e r s e side of colony: No distinctive pigment; yellow-brown on yeast-malt a g a r , light grayish yellow on oatmeal a g a r , s a l t s - s t a r c h a g d r and glycerol-asparagine a g a r . Color i n medium: Melanoid pigment i s not f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment i s found i n the medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r g lycero 1-asparagine agar. Carbon utilization: 2-Glucose, &-arabinose, _D-xylose, g-mannitol, D-fructose and rhamnose a r e utilized f o r growth. No growth o r only t r a c e of growth on raffinose. Utilization of s u c r o s e a n d i - i n o s i t o l is doubtful.
Streptomyces bellus Margalith and B e r e t t a . Description: Margalith and B e r e t t a 1960, 189-195. Type strain: A/870 ( x d . ) . ISP 5185 f r o m L.G. Silvestri, Lepetit S.p.A. a s Lep. M. A/870. ISP description by Group A-13. Spore chain morphology: Section Spirates. Short s p o r e chains may f o r m incomplete spirals, hooks, and loops of s m a l l d i a m e t e r o r flexuous chains (Fig. 30). S p i r a l s a r e best developed on s a l t s - s t a r c h a g a r . Hooks and loops a r e of s m a l l d i a m e t e r and therefore a r e not typical of RA cultures. Mature spore chains with 3 to LO, o r sometimes m o r e than LO, s p o r e s p e r chain on yeast-malt a g a r , oatmeal a e a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Spiny (Fig. 31). Color of colony: A e r i a l m a s s color i n the Blue c o l o r - s e r i e s on oatm e a l a g a r a n d s a l t s - s t a r c h a g a r ; White o r Blue c o l o r - s e r i e s on yeastm a t t a g a r and glycerol-asparagine a g a r . R e v e r s e side of colony: Substrate color is modified by r e d (orange) pigment on yeast-malt a g a r and glycerol-asparagine a g a r and by r e d o r blue pigments on oatmeal a g a r a n d s a l t s - s t a r c h a g a r . These pigments a r e not pH indicators. Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r and tryptone-yeast broth. T r a c e of r e d pigment m a y be found i n the medium i n glycerol-asparagine a g a r ; it i s not a pH indicator. Pigments a r e not found i n the medium i n yeast-malt a g a r , oatmeal a g a r o r s a l t s - s t a r c h a g a r . Carbon uti t i za tion: _D -G lucos e, k-ara binos e, s u c r o se, E-xy Lose i-inositot, 2-mannitol, g.fructose, rhamnose and raffinose are utilized f o r growth.
300
INTERNATIONAL
JOURNAL
Streptomyces bikiniensis Johnstone a n d Waksman. Descriptions: Johnstone a n d Waksman 1947, 294; Johnstone and Waksman 1948, 317326. Type strain: IMRU 3514 (S.A. Waksman a n d R. Gordon, p e r s o n a l communication). ISP 5235 f r o m S.A. Waksman a s IMRU 3514. ISP description by C r o u p B-9. Spore chain morphology: Section Rectiflexibiles (Fig. 32). Mature s p o r e chains LO to 50 o r often m o r e than 50 s p o r e s p e r chain. This morphology i s s e e n on y e a s t - m a l t a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r , and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 33). Color of colony: A e r i a l m a s s color i n the Yellow color-series on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d g l y c e r o l - a s p a r agine a g a r . R e v e r s e side of colony: No distinctive pigment (grayed yellow to grayed yellowish brown) on y e a s t - m a t t a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Color i n medium: Melanoid pigment i s not produced i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. T r a c e of yellow pigment, o r no pigment may be found in medium i n tryptone-yeast broth, y e a s t - m a l t a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r agine a g a r ; this pigment is not a pH indicator. Carbon utilization: _D-GLucose, _D-xylose, D_-mannitol a n d _D-fructose a r e utilized f o r growth. N o growth o r only t r a c e of growth on rhamnose. Utilization of _L-arabinose, s u c r o s e , i - i n o s i t o l and raffinose if doubtful. Actinomyces biverticiltatus Preobrazhenskaya. Description: Preobrazhenskaya i a Gauze e t a t . 1957, 75-76. Type s t r a i n : 10204/54 (G. F. Gauze, p e r s o n a l communication; C. F. Gauze cited by Cottlieb, 1968, 20). ISP 5272 f r o m G. F. Gauze a s 10204/54. Streptomyces biverticillatus (Preobrazhenskaya) Pridham gal,, 1958, 72; Strepto v e r t i c i l l i u m biverticillatus (sic) (Preobrazhenskaya) Baldacci 1958,25; Streptoverticillium biverticillatum (Preobrazhenskaya) B a ldacci, Farina and Locci 1966, 157. The original a u t h o r s now consider this to be identical with Streptomyces h i r o s h i m e n s i s Shinobu 1955, personal communication, 1964. ISP description by Group B -3. Spore chain morphology: Section Verticillati with umbellate-monov e r t i c i l l a t e (biverticillate) s p o r e chains (Fig. 34). Mature s p o r e chains generally contain 10 o r m o r e s p o r e s p e r chain. This morphology i s s e e n on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r agine a g a r . Spore surface: Smooth (Fig. 35). Color of colony: A e r i a l m a s s color i n the Red c o l o r - s e r i e s on y e a s t m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . (One o b s e r v e r , only, placed this culture i n the Violet c o l o r s e r i e s on y e a s t - m a l t a g a r a n d s a l t s - s t a r c h agar). R e v e r s e side of colony: R e v e r s e color i s r e d on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ; this pigment i s pH sensitive, changing f r o m r e d t o o r a n g e - r e d with addition of 0.05 N NaOH a n d f r o m r e d t o violet-red o r blue with addition of 0.05 N HC1. Color i n medium: Melanoid pigments a r e f o r m e d i n l e s s than 2 days i n peptone-yeast-iron a g a r and tryptone-yeast broth, but m o r e slowly i n tyrosine a g a r . P i g m e n t s other than melanoids a r e not f o r m e d i n y e a s t m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine agar.
F i g u r e 3 0 . S. bellus. (a) S p i r a l s p o r e chains on s a l t s - s t a r c h a g a r , 21 d a y s . 23-(b)mplete s p i r a l s , hooks and loops ( X 550) on s a l t s s t a r c h a g a r , 24 d a y s . l 5 F i g u r e 31. S.-bellus. Spiny s p o r e s , c h r o m i u m shadowed; e l e c t r o n m i c r o g r a p h f r o m 24 day c u l t u r e on y e a s t - m a l t a g a r . 24 F i g u r e 3 2 . S. bikiniensis. R F s p o r e chains ( X 240) on y e a s t - m a l t a g a r , 14 days: 2 5 F i g u r e 3 3 . S. bikiniensis. Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 1 4 day c u l t u r e o n y e a s t -malt a g a r . F i g u r e 34. A. b i v e r t i c i l l a t u s . BIV (umbellate monoverticillate) morphoiogy ( X 6 0 0 ) . (a) O n y e a s t - m a l t a g a r , 7 days; (b) On g l y c e r o l a s p a r a g i n e a g a r , 14 days. 2o F i g u r e 35. 4. b i v e r t i c i l l a t u s . Smooth s p o r e s ; Blectron m i c r o g r a p h f r o m 1 4 day c u l t u r e on s a l t s - s t a r c h a g a r . l o
'
302
INTERNATIONAL
JOURNAL
Carbon utilization: _D-Glucose and possibly i - i n o s i t o l and _D-fructose a r e utilized f o r growth; growth o n i - i n o s i t o l a n d 2 - f r u c t o s e i s much l e s s abundant than on _D- glucose. No growth o r only t r a c e of growth on L-arabinose, s u c r o s e , _D-xylose, _D-mannitol, r hamnose and raffinose. Streptomyces bobili (Waksman and C u r t i s ) Waksman and Henrici. Descriptions: Actinomyces bobili Waksman and C u r t i s 1916, 121 a n d 127; Waksman 1919, 100-102; Streptomyces bobiliae (&)(Waksman and C u r t i s ) Waksman a n d Henrici 1948, 937. Type s t r a i n : IMRU 3310 (Waksman 1961, 182). ISP 5056 f r o m S.A. Waksman a s I U R U 3310. ISP d e s c r i p tion by Group B -11. Spore chain morphology: Section Spirales. Sporulating a e r i a l mycelium is not produced a t 27'c o r 3 7 ' C on y e a s t - m a l t agar, oatmeal a g a r o r glycerol-asparagine a g a r , o r on supplementary media (Czapekls a g a r and potato-glucose a g a r ) used by one o b s e r v e r . Two o b s e r v e r s found some spots of sporulating a e r i a l mycelium on s a l t s - s t a r c h a g a r . One collaborator (J.B. Routien) found s p i r a l s p o r e chains with LO to 30 o r m o r e s p o r e s p e r chain on P r i d h a m and Gottliebls b a s a l s a l t s medium f o r carbon utilization enriched with xylose, raffinose o r glucose (Fig. 36). Incubation was a t 37C. S p i r a l s w e r e m o s t abundant when xylose was the carbon s o u r c e . The f i r s t description by Waksman (1916 9. cit.) notes the absence of t r u e a e r i a l mycelium o r s p o r e s on Czapek's medium. The description published i n 1919 (Waksman, 9. notes s p i r a l f o r m a tion on scant white a e r i a l mycelium on glycerin-synthetic a g a r and t r a c e s of a e r i a l mycelium in spots only on s t a r c h a g a r plate and potato plug. A e r i a l mycelium was not produced on 7 other solid media used by Waksman. The p r e s e n t culture s e e m s to conform well to the e a r l y descriptions. Spore s u r f a c e : Smooth (Fig. 37). Color of colony: A e r i a l m a s s color cannot be observed on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r , S p a r s e a e r i a l mycelium, when produced, is white. R e v e r s e side of colony: Substrate mycelium color may be g r a y i s h yellow, o r may be modified with r e d to r e d d i s h gray o r pink, reddish brown, o r reddish orange. The s u b s t r a t e mycelium color i s dependent on pH, changing f r o m yellowish pink to violet pink with addition of 0.05 N NaOH and f r o m yellowish pink to yellow orange with 0.05 N HCl Color i n medium: Melanoid pigments a r e not produced i n peptoney e a s t - i r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. N o pigments a r e found i n medium i n yeast-matt a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-Glucose, _L-arabinose, s u c r o s e , _D-xytose, i -inositol, 2 - f r u c t o s e , r h a m n o s e a n d raffinose a r e utilized f o r growth. N o growth o r only t r a c e of growth on 2-mannitol.
e.)
Streptomyces calvus Backus, T r e s n e r and Campbell. Description: Backus, T r e s n e r and Campbell 1957, 532-541. Type s t r a i n : T-3018 ISP 5010 f r o m H. T r e s n e r a s T-3018. ISP description by Group B-LO. Spore chain morphology: Section S p i r a l e s on s a l t s - s t a r c h a g a r . (Fig. 38). Spore chains a r e s h o r t and poorly developed or absent on y e a s t - m a l t a g a r , oatmeal a g a r and glycerol-asparagine a g a r . The original description by Backus, T r e s n e r and Campbell (%. C t . ) a l s o indicates that a e r i a l mycelium i s poorly developed on m o s t media with
w.).
F i g u r e 3 6 . S. bobili. S p i r a l s p o r e chains ( X 50) on P r i d h a m and Gottlieb carbonGti1ization medium plus raffinose. 36 S._ bobili. Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h ( X 2, 700) F i g u r e 37. _ . f r o m 1 3 day c u l t u r e on P r i d h a m and Gottlieb c a r b o n utilization m e d i u m plus raffinose. 36 Figure 38. S. calvus. S p i r a l s p o r e chains o n s a l t s - s t a r c h a g a r . l 6 F i g u r e 3 9 . S . calvus. Spiny to h a i r y s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 21 day F u l h G n s a l t s - s t a r c h a g a r . F i g u r e 40. A. candidus. R F s p o r e chains ( X 8 7 5 ) . (a) Flexuous a e r i a l chains on y e a s t - m a l t agar, 22 days; (b) Long chains on glycerola s p a r a g i n e a g a r s u r f a c e , 22 d a y s . 27 F i g u r e 41. A . -candidus. Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 14 day c u l t u r e o n o a t m e a l a g a r . l 2
304
INTERNATIONAL
JOURNAL
best sporutation and s p i r a l development on s t a r c h containing inorganic s a l t s media. Spore surface: Spiny to hairy (Fig. 39). Color of colony: A e r i a l mycelium i s inadequate f o r color d e t e r m i n a tion on m o s t media. When m a t u r e sporulating a e r i a l mycelium i s f o r m e d on y e a s t - m a t t a g a r o r s a l t s - s t a r c h a g a r , i t is i n the G r a y c o l o r - s e r i e s . R e v e r s e side of colony: No distinctive pigments (colorless o r g r a y i s h yettowish brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r o r glycerol-asparagine a g a r . Color in medium: Melanoid pigments a r e not found i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth; no pigment is found i n medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerolasparagine agar. Carbon utilization: _D-GLucose, _L-arabinose, s u c r o s e , _D-xylose, - -inositol, _D-mannitol, 2 - f r u c t o s e , rhamnose a n d raffinose a r e a l l i utilized f o r growth. Actinomyce s candidus Krasil'nikov. Descriptions : K r a s i llnikov 1941, 49; Krasil'nikov 1949, LOO. Type s t r a i n : 5855/54 (neotype, P r e o b r a zhenskaya, persona 1 communication ; K r a sill nikov, personal communication). Streptomyces candidus (Krasil'nikov) Waksman 195 3, 94. ISP 5141 f r o m T . P . Preobrazhenakaya a s 5855/54. ISP d e s c r i p tion by Group A-3. Spore chain morphology: Section Rectiflexibiles. Mature s p o r e chains generally long and flexuous, often with m o r e than 50 s p o r e s p e r chain (Fig. 40). This morphology i s seen on s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . Sporulation may be poor on y e a s t - m a l t a g a r and oatmeat a g a r . Spore surface: Smooth (Fig. 41). Color of colony: A e r i a l m a s s color in the White c o l o r - s e r i e s on y e a s t - m a l t a g a r , oatmeat a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r agine a g a r , although a e r i a l mycelium may be poorly developed on y e a s t maL t a g a r and oatmea 1 a g a r . R e v e r s e side of colony: No distinctive pigments ( c o l o r l e s s o r v e r y pate grayed yellow) on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Color i n medium: Melanoid pigments not f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeaat broth. P i g m e n t s a r e not f o r m e d i n medium in y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r g ly ce r o 1-a sparagine a g a r . Carbon uti Lization: ,D -G tucose, _L-arabinose, ._D -xylose, _D -mannit01 and rhamnose a r e utilized f o r growth. N o growth o r only t r a c e of growth on s u c r o s e , i - i n o s i t o l and raffinose.
-e t at.
Streptomyces capreolus Higgens. Description: Higgens & S t a r k 1962, 596-606. Type s t r a i n : M48-E2655 = NRRL 2773 (D.H. Lively, Lilly Res. Labs., personal communication). ISP 5225 f r o m D.H. Lively a s M48-EZ655 = NRRL 2773. ISP description by Group B-2. Spore chain morphology: Section Rectiflexibiles with long s p o r e chains of 50 o r m o r e s p o r e s p e r chain on m a t u r e c u l t u r e s (Fig. 42) on y e a s t malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 43). Color of colony: A e r i a l m a s s color i s usually in the Red o r Yellow c o l o r - s e r i e s a f t e r 14 to 21 days. I m m a t u r e a e r i a l mycelium i s in the White c o l o r - s e r i e s . Color tabs selected by o b s e r v e r s f o r 21 day c u l t u r e s
SYSTEMATIC
BACTERIOLOGY
305
included 2ba (pale yellow) f r o m the Yellow c o l o r - s e r i e s and 3ca (pale orange yellow) to 5cb (grayish yellowish pink) f r o m the Red colors e r i e s . These colors a r e observed on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . R e v e r s e side of colony: Grayed yellow to orange yellow o r orange on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerolasparagine a g a r ; this pigment is not a pH indicator. Color i n medium: Melanoid pigments a r e not f o r m e d in peptoneyeast-iron a g a r , tyrosine a g a r o r tryptone-yeast broth. Yellow o r yellow orange pigment i s found in the medium i n yeast-malt a g a r , oatm e a l a g a r , s a l t s - s t a r c h a g a r and usually glycerol-asparagine a g a r ; this pigment i s not a pH indicator. Carbon utilization: _D-GLucose, _L-arabinose and i - i n o s i t o l a r e utilized f o r growth. Reports v a r y on utilization of _D-xylose, _D-mannitol and D-fructose, although i n each case, two o r t h r e e o b s e r v e r s reported utilization. A t r a c e of growth i s found on the carbon f r e e control and on sucrose, rhamnose and raffinose; these s u g a r s a r e not utilized. Streptomyces catenulae (s1c) Davisson and Finlay. Description: Davisson and Finlay 1959, U . S . Patent 2,895,876; s e e a l s o Waksman 1961, 190-191. Type strain: ATCC 12,476 ( K d . ) = P f i z e r 6563 (J. Routien, Chas. P f i z e r and Co., personal communication). ISP 5258 f r o m J . Routien a s 6563 = ATCC 12,476. ISP description by Group B-5. Spore chain morphology: Section Rectiflexibiles. Very s h o r t spore chains of 3 to LO s p o r e s occur in dense c l u s t e r s s o that morphology i s difficult to determine (Fig. 44a). Short chains often f o r m hooks o r incomplete s p i r a l s (Fig. 44b). Tight s p i r a l s may occur in the dense c l u s t e r s but m o s t chains a r e too short to f o r m t r u e s p i r a l s (Fig. 44c). cultures. This morphology i s Short chains a r e a l s o not typical of seen on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerolasparagine a g a r . It i s representative of morphology described by Davisson and Finlay (%.At.).Spore surface: Smooth (Fig. 45). Color of colony: A e r i a l m a s s color i n the Green c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r and glycerol-asparagine a g a r ; G r a y o r G r e e n c o l o r - s e r i e s on s a l t s - s t a r c h a g a r . The color on a l l media i s medium gray to olive g r a y o r grayish olive (see tabs 2ih o r 2ge i n G r a y c o l o r - s e r i e s and 1 1/2ge i n Green c o l o r - s e r i e s ) . R e v e r s e side of colony: Colorless o r grayish yellow to light grayish olive o r olive brown on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Substrate mycelium pigment i s not pH sensitive. Color in medium: Melanoid pigments a r e not found in peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth; no pigment o r only t r a c e of yellow pigment i n medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Carbon ut lization: _D -Glucose, _D -mannito 1, _D -fructose and raffino s e a r e utilized f o r growth. No growth o r only t r a c e of growth on _L-arabinose, s u c r o s e , _D-xylose, i-inositol and rhamnose. Streptomyce s ce 1l o staticu s Hamada Description: Hamada 1958, ISP 5189 f r o m N. Ishida, Tohoku 173-179. Type strain: E-150 Univ. a s E-150. ISP description by Group A-13.
w.).
F i g u r e 42. S. c a p r e o l u s . Long R F s p o r e chains ( X 380) on Waksman s t a r c h :gar B, 21 days." F i g u r e 4 3 . S. c a p r e o l u s . Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h from 2 1 day c u l t u r e on s a l t s - s t a r c h a g a r . 2 9 F i g u r e 44. S. catenulae. ( a ) Large, d e n s e c l u s t e r s of s p o r e chains ( X 400);n m r c h a g a r , 1 4 days. ( b ) C l u s t e r s of s h o r t s p o r e chains, s o m e of which a p p e a r to be bent into hooks o r p a r t i a l s p i r a l s ( X 400) on s a l t s - s t a r c h a g a r , 14 d a y s . ( c ) Short, t w i s t e d s p o r e chains ( X 300) on s a l t s - s t a r c h a g a r , 14 d a y s . 4 F i g u r e 45. -~ S. catenulae. Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 32 day c u l t u r e on y e a s t - m a l t a g a r . 30
SYSTEMATIC BACTERIOLOGY
307
Spore chain morphology: Section Spirales (Fig. 46). Mature s p o r e chains LO to 50 s p o r e s p e r chain; longer chains a r e often observed. This morphology i s s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Spiny (Fig. 47). Color of colony: A e r i a l m a s s color in the Red (or Gray) colors e r i e s on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerolasparagine a g a r . C h a r a c t e r i s t i c color i s usually between 3ge (light grayish yellowish brown) and 5ge o r 5dc (grayish yellowish pink) color tabs of T r e s n e r - B a c k u s color wheels. R e v e r s e side of colony: N o distinctive pigment ( c h a r a c t e r i s t i c grayed yellow o r light yellow brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r o r glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e f o r m e d in peptone-yeasti r o n a g a r , tyrosine a g a r and tryptone-yeast broth. Pigments other than melanoids o r faint t r a c e s of yellow pigment not formed in yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: 2-Glucose, &-arabinose, sucrose, _D-xylose, i -inositol, D_-mannitol, 2 - f r u c t o s e , rhamnose and raffinose a r e utilized f o r growth. Streptomyces chibaensis Suzuki, Nakamura, Okuma and Tomiyama. Description: Suzuki et a t . 1958, 81-83. Type strain: 77-SN-2 (Suzuki, personal communication). ISP 5220 f r o m S. Suzuki a s 77-SN-2. I S P description by Group B -4. Spore chain morphology: Section Spirales o r Rectiflexibiles. Spirals, when formed, a r e open and poorly developed (Fig. 48a). Some spore chains a r e straight and many a r e flexuous o r curved to f o r m hooks o r p a r t i a l s p i r a l s (Fig: 48b). Moat s p o r e chains a r e too s h o r t (3 to LO o r 20 s p o r e s p e r chain) f o r this culture to be placed in section (Fig. 48b). This morphology is seen on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 49). Color of colony: Mature a e r i a l m a s s color is usually in the G r a y c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r ; G r a y o r Yellow c o l o r - s e r i e s on glycerol-asparagine a g a r . Color tabs selected f r o m the G r a y c o l o r - s e r i e s w e r e yellowish g r a y o r grayish yellowish brown. R e v e r s e side of colony: No distinctive pigments (grayed greenish yellow, yellow o r r a r e l y orange-yettow) on yeast-matt a g a r , oatmeat a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ; substrate mycelium pigment is not a pH indicator. Color i n medium: Melanoid pigments a r e not found in peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. Yellow pigment may o r may not be found in the medium in yeast-malt a g a r , oatmeal a g a r and glycerol-asparagine a g a r ; this pigment i s not pH sensitive. Carbon utilization: _D-GLucose, _L-arabinose, s u c r o s e , D_-xylose, i -inositot, 2-mannitol, _D-fructose, rhamnose and raffinosc a r e a l l utilized f o r growth. Actinomyces chromofuscus Preobrazhenskaya, Ryabova and B linov. Description: Preobrazhenskaya et at. &Gauze et a t . 1957, 176-177. Type strain: 13638/58 (G. F. Gauze, personal communication; G. F.
F i g u r e 46. S . c e l l o s t a t i c u s . S p i r a l s p o r e chains forming tight c o i l s . ( a ) On y e a s t - m a l t a g a r , 21 d a y s . 2 3 (b) On s a l t s - s t a r c h a g a r , 2 2 d a y s ( X 500). l 5 F i g u r e 47. S. c e l l o s t a t i c u s . Spiny s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 13 day c u l t u r e on G a u z e ' s medium N o . 1. F i g u r e 48. S. chibaensis. S p o r e chains ( X 800) o n o a t m e a l a g a r , 14 d a y s : (a) S p i r a l chains; (b) Flexuous chains." F i g u r e 49. S . chibaensis. Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h from 2 1 day c u l t u r e on o a t m e a l a g a r . "
SYSTEMATIC
BACTERIOLOGY
309
Gauze cited by Gottlieb 1968, 20). Streptomyces chromofuscus (Preobrazhenskaya, Ryabova and B linov) P r i d h a m e t at. 1958, 68. ISP 5273 f r o m G.F. Gauze a s 13638/58. ISP Description by Group B-4. Spore chain morphology: Section S p i r a l e s (Fig. 50). Mature s p o r e chains moderately Long with LO to 50, o r often m o r e than 50, s p o r e s p e r chain. This morphology i s s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . Spore surface: Spiny (Fig. 51). Color of colony: A e r i a l m a s s color i n the G r a y c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . R e v e r s e side of colony: No distinctive pigment (grayed yellow to olive brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d g lyce r o 1-a sparagine a g a r . Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r and tryptone-yeast broth, but not i n tyrosine a g a r . Pigments other than melanoids o r faint t r a c e s of yellow a r e not found i n the medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon uti lization: 2-G lucose, _L-arabino s e, ,D --lo se, i - i n o sitol, D-mannitol, _D-fructose and rhamnose a r e utilized f o r growth. Reports v a r y on utilization of raffinose a n d i - i n o s i t o l . NOgrowth o r only t r a c e of growth on sucrose.
Streptoverticillium cinnamoneum (Benedict, Dvonch, Shotwe 11, P r i d h a m and Lindenfelser) Baldacci, F a r i n a , Locci. Streptomyces cinnamoneus Benedict et at. L952, 591-594; Dvonch e t at. 1954,1135-1142; Streptomyces cinnamomeus P r i d h a m e t at. 1956, 576 (orthographic variant); Streptoverticiltium cinnamoneus (81c) (Benedict e t at. ) Baldacci 1958, 25; ,SJreptoverticillium cinnamoneum (Benedict et at. ) emend. hobis Baldacci, F a r i n a , Locci 1966, 158. Type strain: NRRL B-1285 (T. Pridham, p e r s o n a l communication). ISP 5005 f r o m T. P r i d h a m a s NRRL B-1285. ISP description by Group B-1. Spore chain morphology: Umbellate Monoverticillate ( = Streptomyces section Verticillati, biverticillate) (Fig. 52). Mature s p o r e chains generally 3 to LO, o r often m o r e than LO, s p o r e s p e r chain. This m o r phology i s s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 53). Color of colony: A e r i a l mass c o l a i n the Red c o l o r - s e r i e s (grayish yellowish pink) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . R e v e r s e side of colony: No distinctive pigment (grayish yellow to yellow-brown o r brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r . Color i n medium: Melanoid pigments are not f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment i s found i n the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r g Lycerol-asparagine a g a r . Carbon utilization: Vegetative growth i s generally poor on Pridham and Gottlieb's carbon utilization medium plus _D- glucose. Reports f r o m collaborators v a r i e d f r o m no growth to slight growth with other carbon s o u r c e s , but good growth was not observed on any of the following: L-arabinose, s u c r o s e , _D-xylose, g-inositol, _D-mannitol, 2 - f r u c t o s e ,
--
310
F i g u r e 50. A. c h r o m o f u s c u s . S p i r a l s p o r e chains. ( X 400) on o a t m e a l a g a r , 18 d a y s . F i g u r e 51. A . c h r o m o f u s c u s . Spiny s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 14 day c u l t u r e on o a t m e a l a g a r . 31 F i g u r e 52. S t r e p t o v e r t i c i l l i u m cinnamoneum. K V ( u m b e l l a t e m o n o v e r t i c i l l a t e ) s p o r e chains (X 1000) on o a t m e a l a g a r , 14 days. F i g u r e 53. S t r e p t o v e r t i c i l l i u m cinnamoneum. Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 1 4 day c u l t u r e on y e a s t - m a l t a g a r .
SYSTEMATIC
BACTERIOLOGY
311
D- glucose may inaicate a r e q u i r e m e n t f o r a growth factor

i n the b a s a l medium,
rharnnose o r raffinose.
P o o r growth on a l l carbon s o u r c e s including not p r e s e n t
Actinomyces citreof luorescens Korenyako,' Krasil'nikov, Nikitina a n d Sokolova. Description: Korenyako e t at. 1960, 133-159. Type strain: ISP 5265 f r o m N.A. Krasil'nikov a s USSR-RIA 648 = INMI 2292 INMI 2292. ISP description by Group B-2. Spore chain morphology: Section Rectiflexibiles. Mature s p o r e chains long, often with m o r e than 50 s p o r e s p e r chain (Fig. 54). This morphology i s s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 55). Color of colony: A e r i a l m a s s color i n the Yellow c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . R e v e r s e side of colony: No distinctive pigments (light yellow o r g r a y i s h yellow to light olive brown) on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e not produced i n peptoneyeast-iron a g a r , tyrosine a g a r o r tryptone-yeast broth. Yellow pigment i s found i n medium i n glycerol-asparagine a g a r and yellow o r g r e e n pigment i s found i n medium i n yeast-malt a g a r , oatmeal a g a r and s a l t s s t a r c h a g a r . This pigment is not a pH indicator. Carbon utilization: _D-Glucose, &-arabinose, _D-xylose, _D-mannitol, D-fructose a n d rhamnose a r e utilized f o r growth. No growth o r only t r a c e of growth on sucrose, 1-inositol o r raffinose.
m.).
Streptomyces coelicolor (Mutter) Kutzner and Waksman. Descriptions: Mtlller 1908, 197-213; Kutzner and Waksman 1959, 528-538. Not Actinornyces coelicolor (MUller) Lieske 1921, 28; not Streptomyces coelicolor (Mutter) Waksman and Henrici 1948, 935; not Actinomyces violaceus Waksman and C u r t i s 1916, 110 (&ee Kutzner and Waksman 1959, 528-538). Type strain: CBS-R. MII'ller s t r a i n (C.A. deVries, CBS personal communication). ISP 5233 f r o m G.A. d e V r i e s a s CBS-R MUller strain. ISP description by Group B - 2 . Spore chain morphology: Section Rectiflexibiles. Mature s p o r e chains usually 10 to 50, o r often m o r e than 50, s p o r e s p e r chain. (Fig. 56). This morphology i s seen on yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 57). Special morphological c h a r a c t e r i s t i c s : One o b s e r v e r r e p o r t s subs t r a t e conidia on oatmeal a g a r , E m e r s o n ' s potato dextrose a g a r and potato c a r r o t a g a r . Sclerotia formation on yeast-malt a g a r a n d oatmeal a g a r was r e p o r t e d by one o b s e r v e r . Color i n colony: A e r i a l m a s s color i n the Yellow c o l o r - s e r i e s on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . R e v e r s e side of colony: Light olive brown to strong brown o r v e r y d a r k brown on yeast-malt a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r ; brown s u b s t r a t e mycelium pigment on oatmeal a g a r i s pH sensitive changing f r o m brown to g r e e n with addition of 0.05 N NaOH. Potato plug i s pigmented d a r k blue i n 4-7 days.
312
Color in medium: Melanoid pigments a r e not f o r m e d i n peptoneyeast-iron a g a r , tyrosine a g a r o r tryptone-yeast broth. Reddish, yellowish brown , o r g r e e n i s h pigments a r e found i n the medium i n yeast-matt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r agine a g a r ; yellowish brown pigment is changed to g r e e n with addition of 0.05 N NaOH. Liquid surrounding d a r k blue potato plug is pigmented greenish brown in 4-7 days. Carbon utilization: _D-Clucose, I=-arabinose, Q-xylose, _D-mannitol and _D-fructose a r e utilized f o r growth. No growth o r only t r a c e of growth on s u c r o s e , L-inositol, rhamnose and raffinose. Actinomyce s coeruleorubidus P r e o b r a zhenskaya. Description: Preobrazhenskaya k G a u z e e t a t . 1957, 123 and 125. Type strain: Gottlieb 1968). Streptomyces 12531/54 (Ibrd. ; s e e a l s o Gauze coeruleorubidus (Preobrazhenskaya) P r i d h a m e t a l . 1958, 67. ISP 5145 f r o m T. P. Preobrazhenskaya a s 12531/54. ISP description by Group A - 1 3 . Spore chain morphology: Section Spirales. S a l t s - s t a r c h a g a r shows b e s t s p i r a l morphology (Fig. 58a); incomplete s p i r a l s and hooks may a l s o be common, especially on y e a s t - m a l t a g a r , oatmeal a g a r and glycerol-asparagine a g a r (Fig. 58b). Mature s p o r e chains generally 10 to 50 s p o r e s p e r chain. Spore surface: Spiny (Fig. 59). Color of colony: A e r i a l m a s s color i n the Blue c o l o r - s e r i e s on oatmeat a g a r and s a l t s - s t a r c h a g a r ; Blue o r White c o l o r - s e r i e s on yeast-matt a g a r and glycerol-asparagine a g a r . R e v e r s e side of colony: G r a y i s h yellow to yellowish brown on y e a s t m a l t a g a r ; grayed yellow to yellowish g r e e n on oatmeal a g a r and s a l t s s t a r c h a g a r ; both grayed yellowish g r e e n and grayed r e d o r orange on glycerol-asparagine a g a r . These s u b s t r a t e mycelium pigments a r e not pH indicators when tested with 0.05 N NaOH o r H C l . Color in medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r a n d tryptone-yeast broth. Red pigment may be found i n medium i n yeast-malt a g a r , s a l t s - s t a r c h a g a r and glycerolaspaxagine a g a r a f t e r 14 d a y s ; i t i s not pH sensitive. lucose, &-arabino se, s u c r o s e , _D -xy lose, Carbon utilization: _D-G i.-inositol, 2mannitol, _D-fmctose, rhamnose and raffinose a r e utilized for growth. Streptomyces c o r a l u s Dietz (unpublished data). Description: Not published (Alma Dietz, Upjohn Go., p e r s o n a l communication, Sept. 1964). Type s t r a i n : UC 2098 = NRRL 2442 (Dietz, personal communication). ISP 5256 f r o m A . Dietz, Upjohn Go. a s UC 2098 = NRRL 2442. ISP description by Group B -2. Spore chain morphology: Section Rectiflexibiles (Fig. 60). Some long s p o r e chains f o r m open loops suggesting morphology. Mature s p o r e chains a r e usually v e r y long, often with m o r e than 50 s p o r e s p e r chain. This morphology i s seen on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 61). Color of colony: Mature a e r i a l m a s s color i s usually i n the Red c o l o r - s e r i e s on s a l t s - s t a r c h a g a r and in the Red, Yellow o r White c o l o r - s e r i e s on yeast-malt a g a r and glycerol-asparagine a g a r . A e r i a l mycelium i s poorly developed on oatmeal a g a r .
F i g u r e 54. _A. citreofluorescens. R F s p o r e chains (X 500) on oatmeal a g a r , 7 days.8 F i g u r e 55. A. citreofluorescens. Smooth s p o r e s ; electron m i c r o g r a p h f r o m 21-day culture on glycerol-asparagine a g a r . 2 9 F i g u r e 56. S . coelicolor. RF s p o r e chains (X 380) on Waksman's s t a r c h agar B . = * coelicolor. Smooth s p o r e s ; electron micrograph f r o m F i g u r e 57. 7 day culture on y e a s t - m a l t a g a r . 2 9 coeruleorubidus. Spiral s p o r e chains (X 500): (a) Coils F i g u r e 58. of many t u r n s on s a l t s - s t a r c h agar, 24 days; (b) S p i r a l s on oatmeal agar, 2 4 d a y s . F i g u r e 59. A . coeruleorubidus. Spiny s p o r e s ; electron m i c r o g r a p h f r o m 18 day culture on oatmeal a g a r .
2.
A.
''
314
INTERNATIONAL
JOURNAL
R e v e r s e side of colony: P a l e o r light grayish yellow to orangeyellow on y e a s t - m a l t a g a r , o a t m e a l a g a r and glycerol-asparagine a g a r ; light yellow t o g r a y i s h r e d on s a l t s - s t a r c h a g a r . Substrate mycelium pigment i s a pH indicator changing f r o m yellow CR yellowish b r o w n t o r e d with addition of 0.05 N NaOH. Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r and tryptone-yeast broth. Red pigment is found i n the medium i n s a l t s - s t a r c h a g a r ; t r a c e s of yellow pigment a r e found i n the medium i n yeast-malt a g a r and oatmeal a g a r . This pigment i s changed f r o m yellow to pink with the addition of 0.05 N NaOH. Carbon utilization: _D-Glucose, _L-arabinose, s u c r o s e , 2-xylose, i-inositol, _D-mannitol, _D-fructose, r h a m n o s e a n d raffinose a r e a l l utilized f o r growth. k t i n o m y c e s cyanoalbus Krasil'nikov a n d Agre. Description: Krasil'nikov and Agre 1960, 273-274. Type s t r a i n INMX-414 (ibld., Krasil'nikov p e r s o n a l communication). ISP 5198 f r o m N.A. KraSil'nikov a s INMI-414 = RIA 662. ISP Description by Group A-13. Spore chain morphology: Section S p i r a l e s o r Rectiflexibiles. Distinct s p i r a l s a r e r a r e . Moderately s h o r t chains often with only LO to20 s p o r e s may f o r m strongly wavy chains to open s p i r a l s of 1 to 4 t u r n s (Fig. 62). This morphology is s e e n on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d g lycerol-asparagine a g a r . Spore surface: Spiny (Fig. 63). Color of colony: Mature a e r i a l m a s s color (14 to 21 d a y s ) i s i n the G r e e n c o l o r - s e r i e s on y e a s t - m a t t a g a r and s a l t s - s t a r c h a g a r ; Green, G r a y o r White c o l o r - s e r i e s on oatmeal a g a r and glycerol-asparagine a g a r . A e r i a l mycelium i s usually poorly developed on o a t m e a l a g a r . R e v e r s e s i d e of colony: Grayed yellow to g r a y e d yellow-green on y e a s t - m a l t a g a r a n d o a t m e a l a g a r ; g r a y e d yellow-green to blue on s a l t s s t a r c h a g a r a n d glycerol-asparagine a g a r . S u b s t r a t e mycelium pigment i s not a pH indicator. Color i n medium: Melanoid pigments are not f o r m e d i n peptoneyeast-iron a g a r , t y r o s i n e a g a r o r tryptone-yeast broth. No pigment is found i n the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-aeparagine a g a r . Carbon Utilization: _D-Glucose, _L-arabinose, s u c r o s e , _D-xylose, i-inoeitol, _D-mannitol, _D-fructose and r h a m n o s e a r e utilized f o r growth. No growth o r only t r a c e of growth on raffinose. Streptomyces e h i m e n s i s Nakazawa. Description: J a p a n e s e P a t e n t 2125, F e b . 4, 1954; J a p a n e s e P a t e n t 8148, Nov. LO, 1955 ( s e e a l s o Shibata, Honjo, Tokui and Nakazawa 1954, 168 f o r publication of n a m e a n d antibiotic production only). Type strain: 138 = I F 0 3398 (Nakazawa, p e r s o n a l communication; Shibata e t at. 1954, 168). ISP 5253 f r o m K. Nakazawa, Takeda Chem. Ind., Ltd. a s 138 = LF'O 3398. ISP description by Group B-9. Spore chain morphology: Section Verticillati. Mature a e r i a l mycelium is usually umbellate-monoverticillate (biverticillate), but sporulating a e r i a l m y c e l i u m i e v e r y sparBe o r a b s e n t on yeaat-malt agar, oatmeal a g a r , sa Lts e t a r c h a g a r and glyce rot-aeparagiae a g a r . B ive rticillate epore chains can b e s e e n on P r i d h a m a n d Gottlieb carbon utilization medium plus _D-xylose or _L-arabinose and s o m e t i m e s c a n be found on oatmeal a g a r a f t e r 21 daye (Fig. 64). Spore chains are short, usually
F i g u r e 60. S. coralus. E, v e r y long s p o r e chains (X 500) on oatmeal agar, 7-days. ' F i g u r e 61. S . coralus. Smooth s p o r e s ; electron micrograph f r o m 21 day c u l G r e on oatmeal a g a r . 2 9 F i g u r e 62. _A. cyanoalbus. (a) Open s p i r a l s and imperfect s p i r a l s (X 500) on glycerol-asparagine a g a r , 21 days. (b) Flexuous chains on y e a s t - m a l t a g a r , 21 days. 23 Figure 6 3 . cyanoalbus. Spiny s p o r e s ; electron m i c r o g r a p h f r o m 18 day culture on oatmeal a g a r . F i g u r e 64. 2. ehimensis. Z V (umbellate monoverticillate) morphology (X 1100) on oatmeal a g a r , 21 days. F i g u r e 65. S, ehimensis. Smooth s p o r e s ; electron m i c r o g r a p h f r o m 1 0 day culture on yeast-malt a g a r .
A .
316
INTERNATIONAL
JOURNAL
with 3 to 10 s p o r e s p e r chain. Spore surface: Smooth (Fig. 65). Color of colony: A e r i a l m a s s color f o r m a t u r e sporulating a e r i a l mycelium cannot be determined a c c u r a t e l y because of poor sporulation on y e a s t - m a l t a g a r , s a t t s - s t a r c h a g a r o r glycero 1-asparagine a g a r . When sporulating a e r i a l mycelium o c c u r s on oatmeal a g a r , it i s i n the G r a y c o l o r - s e r i e s . (The s p o r e s u r f a c e of poorly sporulating growth i s generally i n the Yellow o r Red c o l o r - s e r i e s on y e a s t - m a l t a g a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r o r s p a r s e a e r i a l mycelium may be white.) R e v e r s e side of colony: Brown t o d a r k brown on y e a s t - m a l t a g a r ; g r a y i s h yellow, orange yellow o r yellowish brown on oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Substrate mycelium pigment i s not a pH indicator. Color in medium: Melanoid pigments a r e produced i n peptone-yeasti r o n a g a r and tryptone-yeast broth, but only weakly o r not a t a l l on tyrosine a g a r . Yellow o r g r a y i s h yellowish brown pigment may be found i n the medium i n y e a s t - m a l t a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . This pigment is not pH sensitive. Carbon utilization: _D-Glucose, &inositol, _D-Mannitol a n d _D-fructose a r e utilized f o r growth. T r a c e s of growth o c c u r on the carbon f r e e control a n d on other ISP carbon s o u r c e s . S t r e p t o m y c e s endue Anderson a n d Cottlieb 1952. Descriptions: Streptomyces 9-20 Gottlieb e t al. 195 1 , 393-394; Streptomyces endus Gotttieb and C a r t e r 1950, U.S. P a t e n t 2 , 7 4 6 , 9 0 2 ; Upjohn Go. 1954, B r i t i s h P a t e n t Spec. 705,622. See also: Anderson and Gottlieb 1952, ISP 5187 f r o m D. 294-308. Type strain: 9-20 = NRRL 2339 ( E d . ) . Gottlieb a s 9-20 = NRRL 2339. ISP description by Group A-13. Spore chain morphology: Section S p i r a l e s with tightly closed s p i r a l s (Fig. 6 6 ) . Individual s p o r e s a r e not easily detached f r o m the s p i r a l chain aad compact coils m a y become hydroscopic (coalesce i n black m a i s t droplets). Spore surface: Warty (Fig. 67). Color of colony: A e r i a l m a s s color i n the G r a y c o l o r - s e r i e s a f t e r 21 d a y s on yeast-malt a g a r , o a t m e a l a g a r a n d s a l t s - s t a r c h a g a r . A e r i a l mycelium on glycerol-asparagine a g a r a n d young a e r i a l mycelium on o t h e r media m a y be white. Older colonies s o m e t i m e s become m o i s t a n d black. R e v e r s e side of colony: No distinctive pigments (grayed yellow to yellowish brown o r g r a y ) on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r a n d glycerol-asparagine a g a r . In o l d e r c u l t u r e s the oubstrate mycelium sometime s b e c o m e s m o i s t and black. S u b s t r a t e color i s not affected by pH change. Color i n medium: Melanoid pigments not f o r m e d i n peptone-yeasti r o n a g a r , tyroeine a g a r o r tryptone-yeast broth. No pigments a r e found i n the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r or glycerol-asparagine a g a r . Carbon utilization: _D-Glucoee, _L-arabinose, _D-xylose, _D-mannitol, D-fructose a n d r h a m n o s e a r e utilized f o r growth. N o growth o r only trace of growth on sucrose, &-inositol and raffinose.
Streptomyces filipinensis Ammann, Gottlieb, Brock, G a r t e r and Whitfield. Description: Ammann e t at. 1955, 559-563. Type strain: ISP 5112 f r o m D. Gottlieb a s N R R L 2437 = 114-8. ISP 114-8 @d.). description by Group B-3.
F i g u r e 66. endus. S p i r a l s p o r e chains; tight s p i r a l s (X 550) on glycerol-asparagine a g a r , 23 days. l5 F i g u r e 67. S. e s . Warty s p o r e s in tight s p i r a l s ; electron m i c r o g r a p h f r o m 13-day culture on Gauze's medium No. 1. F i g u r e 68. S. filipinensis. S p i r a l spore chains. (a) Tight s p i r a l s a t end of s p o r e c h a i n on y e a s t - m a l t a g a r , 15 days; (b) Open coils (X 500) on y e a s t - m a l t a g a r , 11 days. 2o F i g u r e 69. S. filipinensis. Spiny s p o r e s ; electron m i c r o g r a p h f r o m 14 day culture on y e a s t - m a l t a g a r . lo F i g u r e 70. A. finlayi. (a) Flexuous chains, hooks and imperfect s p i r a l s (X 625);n s a l t s - s t a r c h agar, 14 days.22 (b) I r r e g u l a r s p i r a l s on yeast-malt a g a r , 21 days. 23 F i g u r e 71. _A. finlayi. H a i r y s p o r e s . 2 z
1
2. -
318
Spore chain morphology: Section Spirales. Tight s p i r a l s (Fig, 68a) o r open coils, usually of s e v e r a l t u r n s (Fig. 68b), occur a t the end of moderately tong s p o r e chains of LO to 50 o r m o r e s p o r e s p e r chain on y e a s t - m a l t a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r . S p i r a l s may be replaced by morphology on glycerol-asparagine a g a r . Spore surface: Spiny (Fig. 69). Color of colony: A e r i a l m a s s color is i n the G r a y c o l o r - s e r i e s on yeast-matt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . R e v e r s e side of colony: No distinctive pigments (colorless to grayish yellow) on yeast-matt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g 1y c e r o 1-a spa r agine aga r Color i n medium: Melanoid pigments a r e produced i n peptone-yeasti r o n a g a r , tyrosine a g a r and peptone-yeast broth. Pigments other than melanoids a r e not found i n the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , salts-starch agar or glycerol-aspragine agar. Carbon uti li zation: _D -G luco s e, _L-arabinose , s u c r o s e, _D -xy lo s e n j-inositol, 2-mannitol, _D-fructose a n d raffinose a r e utilized f o r growth. No growth o r only t r a c e of growth on rhamnose.
Actinomyces finlayi Szabb, Marton, Buti and Partai. Description: Szab6 gal. 1963, 207-214. Type strain: R-1-30 = FBUA 1869 k d . ) . ISP 5218 f r o m I. Szab6 a s R-1-30= F B U A 1869. I S P description by Group B-12. Spore chain morphology: Section Rectiflexibiles to Spirales. Short s p o r e chains of LO o r m o r e s p o r e s p e r chain a r e generally flexuous, crooked, hooked o r i n i m p e r f e c t s p i r a l s of only one o r two t u r n s (Fig. 70a); longer chains may f o r m open s p i r a l s of t h r e e o r four t u r n s (Fig. 70b). Hooks and loops on s h o r t s p o r e chains a r e not c h a r a c t e r i s t i c of morphology. This morphology is s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ; s p i r a l s a r e b e s t developed on y e a s t - m a l t a g a r and oatmeal a g a r . Spore surface: Hairy (Fig. 71). Color of colony: A e r i a l m a s s color i n the Gray c o l o r - s e r i e s on y e a s t m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . R e v e r s e side of colony: Grayed yellow to yellow-brown modified by g r e e n (grayish yellow to olive brown on y e a s t - m a l t a g a r and oatmeal a g a r ; pale g r e e n i s h yellow o r yellow-green to moderate o r d a r k olive g r e e n on s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r ) . Substrate mycelium pigment is not a pH indicator. Color in medium: Melanoid pigments a r e not produced i n peptoney e a s t - i r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigments a r e found i n the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-Glucose is utilized f o r growth. Significant growth a l s o o c c u r s on _L-arabinose, _D-xylose and rhamnose, although growth on these carbon s o u r c e s is Less than on _D- glucose. Utilization of s u c r o s e i s doubtful. No growth o r only t r a c e of growth o n i - i n o s i t o l , D-mannitol, _D-fructose and raffinose. Actinomyces f luorescens Korenyako, Krasil'nikov, Nikitina and Sokolova. Description: Korenyako e t a l . 1960, 133-159 ( s e e a l s o Engl. T r a n s . , 128-156). Type strain: I N M I 592 ISP 5203 f r o m N.A. Krasil'nikov a s I N M I 592 = RIA 647. ISP Description by Group B-6.
e.).
SYSTEMATIC
BACTERIOLOGY
319
Spore chain morphology: Section Rectiflexibiles (Fig. 72). Mature s p o r e chains generally LO to 50 s p o r e s p e r chain. This morphology i s seen on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerola s p a r a g i n e a g a r . Spore surface: Smooth (Fig. 73). Color of colony: A e r i a l m a s s color i n the Yellow c o l o r - s e r i e s on y e a s t - m a l t a g a r , oatmeai a g a r and s a l t s - s t a r c h a g a r ; White o r Yellow color - s e r i e s on glycerol-asparagine a g a r . R e v e r s e side of colony: No distinctive pigments (colorless to grayish yellow on oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ; grayed orange-yellow to olive brown on y e a s t - m a l t agar). R e v e r s e pigment i s not pH sensitive o r i s changed only slightly f r o m yellow-brown to yellow-green, with addition of 0.05 N NaOH. Color i n medium: Melanoid pigments a r e not produced i n peptoney e a s t - i r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. A s m a l l amount of yellow pigment m a y be found i n the medium i n yeast-malt a g a r , oatm e a l a g a r , s a l t s - s t a r c h a g a r and tryptone-yeast broth. This pigment when p r e s e n t i s only slightly pH sensitive, changing f r o m yellow-brown to yellow-green with addition of 0.05 N NaOH. Carbon uti liza tion: D_ -G luco s e, _L-arabino s e, _D -xy lo se, _D -mannit0 1, D-fructose and rhamnose a r e utilized f o r growth. No growth or only t r a c e of growth on s u c r o s e , L-inositol o r raffinose. The original d e s cription f o r t h i s species ( = . s t a t e s that rhamnose i s not utilized , noting that this i s one difference b e t w e e n k . fluorescens and other s p e c i e s i n the fluorescent group. However, a l l of the 3 ISP o b s e r v e r s r e c o r d e d good growth on rhamnose, equivalent to growth on _D-grlucose.
c.)
Streptomyces f r a g i l i s Anderson, Ehrlich, Sun and Burkholder. Description: Anderson e t a l . 1956, 100-105 and 114-115. Type strain: isolate ISP 5044 f r o m L. Anderson, C 1437 = P - D 04926 = NRRL 2424 R e s . Lab., P a r k e , Davis and Co. a s P - D 04926 = NRRL 2424. ISP description by Group B-2. Spore chain morphology: Section Spirales to Rectiflexibiles. Short s p o r e chains i n many straight to flexuous chains (Fig. 74a), but hooks o r loops of s m a l l d i a m e t e r and incomplete s p i r a l s a r e a l s o common (Fig. 74b). S p i r a l s of two o r m o r e complete t u r n s a r e a l s o present, but a r e not common, Spore chains a r e generally short, but usually contain LO o r m o r e s p o r e s p e r chain. Spore surface: Smooth with some tendency toward warty s u r f a c e on s p o r e s near the t e r m i n a l p a r t of the chain (Fig. 75). Color of colony: A e r i a l m a s s color i n the Red c o l o r - s e r i e s on y e a s t m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . White a e r i a l mycelium may a l s o be p r e s e n t on s a l t s - s t a r c h a g a r and g lycerol-asparagine a g a r . R e v e r s e side of colony: Grayed yellow to yellow-brown modified by r e d (Light to moderate yellow o r orange-yellow on s a l t s - s t a r c h a g a r ; light g r a y i s h reddish brown to d a r k brown on yeast-malt a g a r , oatmeal a g a r and glycerol-asparagine agar). Reddish brown pigment may change to yellow brown with addition of 0.05 'N HC1. Color in medium: Melanoid pigments a r e not produced i n peptoneyeast-iron a g a r , tyrosine a g a r or trypto ne-yeast broth. Medium may r e m a i n unpigmented o r may contain s m a l l amounts of yellow o r r e d brown pigment i n the mdeium i n yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r o r glycerol-asparagine a g a r . This pigment, when present,
w.),
F i g u r e 7 2 . A. f l u o r e s c e n s . R F s p o r e c h a i n s (X 500) on s a l t s - s t a r c h a g a r , 23 days. l5 F i g u r e 73. S. f l u o r e s c e n s . Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 21 day c u l t u r e o n y e a s t - m a l t a g a r . 32 F i g u r e 74. S. f r a g i l i s . ( X 380) on W a k s m a n ' s s t a r c h a g a r B, 21 d a y s : (a) S t r a i g h t t o flexuous c h a i n s ; (b) Loops, incomplete s p i r a l s and s h o r t s p i r a l s on t h e s a m e medium,'* F i g u r e 75. S. f r a g i l i s . Smooth s p o r e s , with s o m e s u r f a c e i r r e g u l a r i t i e s on t e r m i n a l s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 14 day c u l t u r e on yeast-malt agar. 2 9
SYSTEMATIC
BACTERIOLOGY
321
is pH sensitive changing f r o m red-brown to yellow with addition of 0.05
N HC1. Carbon utilization: D_-Glucose, _L-arabinose and 2-xylose a r e utilized f o r growth. Utilization of s u c r o s e is doubtful. No growth o r only t r a c e of growth o n 2 -inositol, _D-mannitol, 2 - f r u c t o s e , rhamnose o r raffinose. Actinomyces fulvoviridis Kuchaeva, Krasillnikov, Skryabin and Taptykova. Description: Kuchaeva e t a l . 1960, 251 (see a l s o Engl. ; type s t r a i n selected t r a n s l . , 222-25L). Type strain: INMI 16-3 (i&. by N.A. Krasillnikov, personal communication). ISP 5210 f r o m N . A . Krasil'nikov a s INMI 16-3 = RIA 660. ISP description by Group B-12. Spore chain morphology: Section Rectiflexibiles (Fig. 76). Mature s p o r e chains moderately s h o r t with LO to 50 s p o r e s p e r chain. This morphology i s s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 77). Color of colony: A e r i a l m a s s color i n the G r a y c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . R e v e r s e side of colony: Yellow to yellow-brown plus t r a c e s of g r e e n (pale yellow, yellow o r grayed greenish yellow to olive brown o r d a r k olive on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerola s p a r a g i n e a g a r ) . R e v e r s e pigment i s not a pH indicator. Color i n medium: Melanoid pigments a r e not produced in peptone yeast-iron a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment o r only faint t r a c e of yellow i s found i n medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-Glucose, _L-arabinose. _D-xylose, 2-mannitol, D-fructose and rhamnose a r e utilized f o r growth; utilization of s u c r o s e is doubtful. No growth o r only t r a c e of growth o n i - i n o s i t o l o r raffinose. Streptomyces g a l b u s F r o m m e r . Description: F r o m m e r 1959, 187206. Not S. g a l b u s Okami and Umezawa 1959. Type strain: Wind 731 & d . ; type selected by W. F r o m m e r , p e r s o n a l communication). ISP 5089 f r o m C. W. Hesseltine a s NRRL B-2283 = Wind 731 f r o m W. F r o m m e r . ISP description by Group B-5. Spore chain morphology: Section Spirales or Retinaculiaperti. S p i r a l s may be open coils of s e v e r a l turns, tight s p i r a l s a t the ends of flexuous s p o r e chains o r i m p e r f e c t s p i r a l s and loops a t the ends of s p o r e chains suggesting =morphology (Fig. 78). Spore chains mode r a t e l y long with LO to 50, o r often m o r e than 50, s p o r e s p e r chain. Spore surface: Smooth to warty. Surface i r r e g u l a r i t i e s suggesting v e r y s m a l l w a r t s a r e c h a r a c t e r i s t i c (Fig. 79). Color of colony: A e r i a l m a s s color i n the Gray c o l o r - s e r i e s on yeast-malt a g a r and oatmeal a g a r and i n the Yellow c o l o r - s e r i e s o r G r a y c o l o r - s e r i e s on s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . G r a y color i s r e p r e s e n t e d by tabs 2fe (medium g r a y ) o r 3fe (light browni s h g r a y ) on a l l media; and I b a o r 2ba (pale yel1ow)or 1 1/2db (pale g r a y i s h yellow) on s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . R e v e r s e side of colony: No distinctive pigments (yellow to yellowbrown o r olive brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h agar and glycerol-asparagine a g a r .
F i g u r e 76. A. fulvoviridis. R F s p o r e chains on y e a s t - m a l t a g a r , 21 days.23 F i g u r e 77. A. fulvoviridis. Smooth s p o r e s ; electron m i c r o g r a p h f r o m 14 day culture on s a l t s - s t a r c h agar." F i g u r e 78. S . galbus. S p i r a l o r R A s p o r e chains; (a) On oatmeal a g a r , 14 day<(X 400). (b) On glycerol-asparagine a g a r , 16 days. 33 S. galbus. Smooth s p o r e s with surface i r r e g u l a r i t i e s Figure 7 9 . suggesting v e r y s m a l l w a r t s ; electron micrograph f r o m 16 day culture on yeast-malt a g a r .
'
SYSTEMATIC
BACTERIOLOGY
323
Color i n medium: Melanoid pigments a r e produced i n peptone-yeasti r o n a g a r and tryptone-yeast broth, but not i n tyrosine a g a r . Yellow soluble pigment i s found in the medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and gtycero 1-asparagine a g a r . This pigment i s not a pH indicator. Carbon utilization: ,D- Glucose, _L-arabinose, _D-xylose, j--inositol, D-mannitol and _D-fructose a r e utilized f o r growth. Only a t r a c e of growth comparable to growth on the carbon-free control i s found on sucrose, rhamnose and raffinose. Streptomyces g a r d n e r i (Waksman 1942) Waksman 1961. Descriptions: Proactinomyces 2. G a r d n e r and Chain 1942, 123-127; Proactinomyces g a r d n e r i Waksman in Waksman, Horning, Welsch and Woodruff 1942, 289-290; Nocardia p a r d n e r i (Waksman) Waksman and Henrici 1948, 914; Streptomyces g a r d n e r i (Waksman) Waksman 1961, 215-216. Type strain: IMRU 3834 (Waksman 1961, 216). ISP 5064 f r o m S.A. Waksman as IMRU 3834. ISP description by Group B -3. Spore chain morphology: Section Rectiftexibiles (Fig. 80). Spore chains moderately long, usually LO to 50, o r often m o r e than 50, s p o r e s p e r chain. This morphology i s seen on yeast-malt a g a r , oatmeal a g a r , s a t t s - s t a r c h a g a r and glycerol-asparagine a g a r , a tthough a e r i a l mycelium m a y be poorly developed on oatmeal a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 81). Color of colony: A e r i a l m a s s color in the G r a y c o l o r - s e r i e s on s a l t s - s t a r c h a g a r i n 14 to 21 days. Mature a e r i a l mycelium may be inadequate f o r determination of a e r i a l m a s s color on yeast-malt a g a r , oatmeal a g a r and glycerol-asparagine a g a r . One o b s e r v e r obtained good sporulating a e r i a l growth i n the Gray c o l o r - s e r i e s on _ . potato a g a r (potato 200 g m ; a g a r 18 gm; t a p water 1 L; pH 6.9-7.1). R e v e r s e side of colony: No distinctive pigment (colorless to grayish yellow o r grayed yellow-brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r , glycerol-asparagine a g a r and potato a g a r . Color i n medium: Melanoid pigments a r e produced i n peptone-yeasti r o n a g a r and tryptone-yeast broth, but not i n tyrosine a g a r (or only i n t r a c e amounts) i n 2 to 4 days. No pigments a r e found i n the medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerot-asparagine agar. Carbon Utilization: _D-GLucose, _L-arabinose, sucrose, ,D-xylose, D-fructose, rhamnose and raffinose a r e utilized for growth. No growth o r only t r a c e of growth o n i - i n o s i t o l and _D-mannitol. :
Streptomyces getaticus (Waksman) Waksman and Henrici. Description: Actinomyces Waksman 1919, 165-166; Streptomyces g e l a t i c u s (Waksman) Waksman and Henrici 1948, 952 (see a l s o Anderson g 1956, 100-115; and HUtter 1967, 319). Type strain: IMRU 3323 (Waksman 1961, 217). ISP 5065 f r o m S . A . Waksman a s IMRU 3323. ISP d e s c r i p tion by Group B -6. Spore chain morphology: Section Rectiflexibiles (Fig. 82). A e r i a l mycelium i s not procuded on m o s t media. Absence of a e r i a l mycelium on m o s t media w a s a l s o noted i n the original description (Waksman 1919 3 . c & . ) which, however, r e c o r d s open s p i r a l s on the scant growth on synthetic a g a r . ISP o b s e r v e r s found wavy s p o r e chains, but no s p i r a l s i n the thin a e r i a l mycelium on yeast-matt a g a r and gtycerol-asparagine
s .
321
INTERNATIONAL
JOURNAL
a g a r . The absence of s p i r a l s h a s a l s o been noted by Anderson e t a l . (1956%. &.), by Waksman i n a later description (1961o~.&t.), a n d by HUtter ( 1 9 6 7 9 . A t . ) . Spore surface: Smooth (Fig. 8 3 ) . Color of colony: A e r i a l m y c e l i u m i s poorly developed on m o s t media. When a n adequate s p o r e m a s s is produced i t i s i n the G r a y c o l o r - s e r i e s . This was observed on yeast-malt a g a r a n d glycerol-asparagine a g a r . Additional media used unsuccessfully by collaborators i n a n effort to get good sporulating growth included Bennett's a g a r , Anderson's a g a r , blood a g a r , Lemco a g a r a n d milk a g a r . R e v e r s e side of colony: No distinctive pigments (colorless to grayish yellow) on y e a s t - m a l t a g a r , oatmeat a g a r , s a l t s - s t a r c h a g a r o r glycerolasparagine agar. Color i n medium: Melanoid pigments a r e not produced i n peptoney e a s t - i r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment i s found i n medium in y e a s t - m a l t agar, oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: Q-Glucose, s u c r o s e , _D-xylose and r h a m n o s e a r e utilized f o r growth. No growth o r only t r a c e of growth on I=-arabinose, i'-inositol, Q m a n n i t o l , Q-fructose and raffinose. Actinomyce s g lobisporus K r a si llnikov. Description: K r a si llnikov 1941, 48. Streptomyces globisporus (Krasil'nikov) Waksman 1953, 39. Type s t r a i n : INMI 2 3 0 2 = RLA 335 ( N A . Krasil'nikov, p e r s o n a l communication). ISP 5199 f r o m N.A. Krasil'nikov a s INMI 2302 = RIA 335. ISP description by Group B -2. Spore chain morphology: Section Rectiflexibiles (Fig. 84). Mature s p o r e chains predominantly flexuous and moderately long with LO to 50, o r often m o r e than 50, s p o r e s p e r chain. This morphology i s seen on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r agine a g a r . Spore surface: Smooth (Fig. 85). Color of colony: A e r i a l m a s s color i n the Yellow c o l o r - s e r i e s on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r agine a g a r . R e v e r s e side of colony: No distinctive pigments (pale yellow to light olive brown) on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glyce rot-a sparagine a g a r . Color in medium: Melanoid pigments a r e not produced i n peptoneyeast-iron a g a r , tyrosine a g a r o r tryptone-yeast broth. Yellow to greenish yellow pigment may be found i n medium in y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . This pigment is not sensitive when tested with 0.05 N HC1 o r NaOH. Carbon utilization: D-Glucose, L-arabinose, D-xylose, D-mannitol, D-fructose and r h a m n o i e a r e utilized for growth, N o grou?h o r only t r a c e of growth on s u c r o s e , i-inositol o r raffinose. Streptomyces goshikiensis Niida. Description: Niida, J a p a n e s e P a t e n t 425909, Japan Official Gazette 38-26948 (Engl. T r a n s l . f r o m T. Niida, p e r s o n a l communication). Name without description: Streptomyces g o s h i k i e m i s Niida Kondo, Sakamoto a n d Yumoto 1961, 365 and i n Kondo e t a l . 1962, 157. Type strain: ISP 5190 received f r o m M. Oda, Meiji Seika Go. a s holotype strain. ISP description by group A-13. Spore chain morphology: Section Spirales, with open s p i r a l s of 4 to
Figure 80. S. gardneri. RF spore chains (X 600) on salts-starch agar,

16
Figure 81. S. gardneri. Smooth spores; electron micrograph from 14 day &tu=yeast-malt agar. lo spore chains on salts-starch agar, 23 days. Figure 82. 2. gelaticus. Figure 83. 2. elaticus. Smooth spores, Au/Pd shadowed; electron micrograp! from 2 1 day culture on oatxheal agar. Figure 84. _A. globisporus. RF spore chains on: (a) Oatmeal agar, 7 days (X 1000). (b) W a k G a n s starch agar B, 2 1 days (X 380). Figure 85. 2. globisporus. Smooth spores; electron micrograph f r o m 14 day culture on oatmeal agar. 2 9
326
INTERNATIONAL
JOURNAL
10 convolutions a t the end of long s p o r e chains (Fig. 86). Mature s p o r e chains LO to 50 s p o r e s p e r chain; longer chains a r e often observed. This morphology i s s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth with m i n o r s u r f a c e i r r e g u l a r i t i e s , but no t r u e spines (Fig. 87). Color of colony: A e r i a l m a s s color i n the Red (or G r a y ) c o l o r - s e r i e s on y e a s t - m a l t a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r . C h a r a c t e r i s t i c color on these media is between 3ge (light g r a y i s h yellowish brown) and 5dc o r 6ec (grayish yellowish pink) color t a b s of T r e s n e r - B a c k u s Color Wheela. A e r i a l mycelium m a y be poorly developed on glycerol-asparagine agar. R e v e r s e side of colony: No distinctive pigments ( c h a r a c t e r i s t i c grayed yellow on oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r to light orange yellow o r light brown on y e a s t - m a l t a g a r ) . C o l a i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r a n d tryptone-yeast broth. P i g m e n t s other than melanoids o r faint t r a c e s of yellow a r e not produced i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Carbon utilization: _D-Glucose a n d possibly _D-fructose a r e utilized f o r growth. Only a t r a c e of growth comparable to growth on carbonf r e e b a s a l medium on L-arabinose, s u c r o s e , _D-xylose, L-inositol, D-rnannitol, rhamnose and raffinose.
St reptomyce s g r i seinus W aksman De scription: G ris ein -producing s t r a i n of Streptomyces g r i s e u s , Reynolds and Waksman 1948, 739-752; Streptomyces g r i s e i n u s Waksman 1959, 1043-1046. Type strain: IMRU 3478 (Waksman 1959, 1045). ISP 5047 f r o m S.A. Waksman a s IMRU 3478. ISP description by Group B-3. Spore chain morphology: Section Rectiflexibles (Fig. 88). Mature s p o r e chains predominantly flexuous a n d moderately long with LO to 50, or often m o r e than 50, s p o r e s p e r chain on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 89). Color of colony: A e r i a l mass color i n the Yellow c o l o r - s e r i e s on s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r a n d i n the Yellow o r G r a y c o l o r - s e r i e s on oatmeal a g a r and yeast-malt a g a r . N e a r e s t matching color tab f r o m the G r a y c o l o r - s e r i e s is Zdc, yellowish gray. R e v e r s e side of colony: No distinctive pigments (grayish yetlow to yellowish brown or light olive brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . The s u b s t r a t e mycelium pigment may be changed slightly f r o m pale yellow to pale yellow-brown o r pale violet with addition of 0 . 0 5 N NaOH, o r f r o m yellowish brown to pale yellowwith 0 . 0 5 N HC1. Color i n medium: Melanoid pigments a r e not produced i n peptoney e a s t - i r o n a g a r , t y r o s i n e a g a r o r tryptone-yeast broth. A t r a c e of yellow to reddish brown pigment may be found i n the medium i n oatmeal a g a r . This pigment, when preeent, is slightly pH sensitive changing f r o m pale yellow to pale violet with addition of 0 . 0 5 N NaOH. Carbon utilization: _D-Glucose, _L-arabinose, _D-xylose, _D-mandtol, D-fructose and r h a m n o s e a re utilized f o r growth. No growth o r only trace o f growth on s u c r o s e , 1-inositol or raffinose.
Streptomyces griaeofuacus Sakamoto, Kondo, Yumoto and Arishima. Deecription: Sakamoto e t at. 1962, 98-101. Type strain- 1068 N d . ) .
Figure 86. S. goshikiensis. Spiral spore chains (X 500) on oatmeal agar, 22 Figure 87. 2. goshikiensis. Smooth spores w i t h minor surface irregularities, but no true spines. chromium shadowed. Electron micrograph f r o m 22 day culture on oatmeal agar. spore chains on yeast-malt agar, 15 days. l1 Figure 88. 2. griseinus. Figure 89. 2. grieeinus. Smooth spores; electron micrograph from 14 day culture on oatmeal agar. lo
*'
328
ISP 5191 f r o m M. Oda, Meiji Seika Co. as 1068. ISP description by Group A-13. Spore chain morphology: Section S p i r a l e s (Fig. 90). Mature s p o r e chains generally LO to 50 s p o r e s p e r chain. This morphology i s observed on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-aspar agine a g a r . Spore surface: Smooth (Fig. 91). Color of colony: A e r i a l m a s s color i n the Red o r G r a y c o l o r - s e r i e s on yeast-malt a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r and glycerolasparagine a g a r . C h a r a c t e r i s t i c color i s between 4ge o r 5ge (light grayish reddish brown) and 4ig (tight g r a y i s h brown) color tabs of T r e s n e r - B a c k u s color wheels. R e v e r s e side of colony: No distinctive pigments on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Color i n medium: Melanoid pigments not f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment found i n medium i n y e a s t - m a l t a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r o r glycerolasparagine agar. Carbon utilization: _D-Glucose, &-arabinose, _D-xylose, 2 - m a n n i t o l and _D-fructose a r e utilized f o r growth. No growth o r only t r a c e of growth on s u c r o s e , i-inositol, rhamnose and raffinose.
Ac tinomyc e s g r i s e oincarnatu s Pr eob ra zhe nskaya , Rya bova and BLinov. Description: Preobrazhenskaya e t at. &Gauze et at. 1957, 1 6 9 171. Type strain: 9673/55 (C. F. Gauze, p e r s o n a l communication: G. F. Gauze, cited by Gottlieb 1968, 2 0 ) . Streptomyces griseoincarnatus (Preobrazhenskaya e t aL.) P r i d h a m et at. 1958, 69. ISP 5 2 7 4 f r o m G.F. Gauze as 9673/55. ISP description by Group B -12. Spore chain morphology: Section S p i r a l e 5 S p i r a l s a r e generally open with 5 to 7 o r m o r e t u r n s (Fig. 92a);flexuous s p o r e chains and i m p e r f e c t s p i r a l s suggesting &morphology a r e a l s o common (Fig. 92b). S p i r a l s a r e b e s t developed on oatmeal a g a r and s a l t s - s t a r c h a g a r . Spore chains a r e moderately long, especially on oatmeal a g a r and s a l t s - s t a r c h a g a r with LO to 5 0 , o r often m o r e than 50, s p o r e s p e r chain. Spore s u r f a c e $piny (Fig. 93). Color of colony: A e r i a l m a s s color i n the G r a y c o l o r - s e r i e s on s a l t s s t a r c h a g a r a f t e r 14-21 d a y s ; younger a e r i a l mycelium may be i n the Red c o l o r - s e r i e s . A e r i a l m a s s color i n the G r a y o r B e d c o l o r - s e r i e s on y e a s t - m a l t a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r . O b s e r v e r s did not a g r e e on the selection of individual color tabs r e p r e s e n t i n g the a e r i a l color, but a l l chose t a b s ranging f r o m grayish yellowish pink o r grayish reddish brown to medium gray. R e v e r s e side of colony: Yellow t o yellow-brown plus r e d (grayish yellow on e a l t s - s t a r c h a g a r ; orange-yellow to reddish brown on y e a s t m a l t a g a r , oatmeal a g a r and glycerol-asparagine a g a r ) , Substrate mycelium pigment i s not a pH indicator. Color i n medium: Melanoid pigments a r e not produced in peptoney e a s t - i r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. Red o r orange pigment i s found i n the medium i n yeast-malt a g a r i n 7 d a y s and i n oatmeal a g a r i n 14-21 days. Red o r orange pigment may o r may not be found i n glycerol-aeparagine a g a r . This pigment is not a pH indicator, Carbon utilization: _D-Glucose, &-arabinose, s u c r o s e , ,D -xylose, _Dmannitol, _D-fructose and rhamnose a r e utilized f o r growth. Reports v a r y on utilization o f i - i n o s i t o l and only a t r a c e of growth i s found on raffinose.
Figure 90. 2. griseofuscus. S iral spore chains (X 500) on glycerolasparagine agar, 22 days. Figyre 9 1 . S . griseofuscus. Smooth spores; electron micrograph from 18 day culture on oatmeal agar. Figure 9 2 . A. griseoincarnatus. Spiral spore c h a b j (a) Opsn @rda on oatmeai-agar, 21 days. (b) Imperfect spirals (X 600) on saltsstarch agar, 14 days. Figure 93. A. griseoincarnatus. Spiny spores; electron micrograph from 14day culture on oatmeal agar. 22
3 30
INT E R N A T I O N A L J O U R N A L
Actinomyces g r i a e o r u b e r Ryabova and Preobrazhenskaya. Description: Ryabova and Preobrazhenskaya in Gauze e t at. 1957, 186 and 195. Streptomyces Rriseoruber (Ryabova and Preobrazhenskaya) P r i d h a m -e t at. 1958, 69. Not Streptomycee Kriseoruber Yamaguchi and Saburi 1955. Type strain: 2022/55 (G. F. Gauze, personal communication 1964; G.F. Gauze&Gottlieb 1968, 2 0 ) . ISP 5275 f r o m C . F . Gauze as 2022/55. ISP description by Group B-5. Spore chain morphology: Section Spirales (Fig. 94). Spore chains moderately long with LO to 50, o r often m o r e than 50, s p o r e s p e r chain. This morphology is seen on yeast-malt a g a r , oatmeal a g a r , salts-starch a g a r and glycerol-asparagine a g a r . Spore surface: Spiny (Fig. 95). Color of colony: Aerial m a s s color in the Gray c o l o r - s e r i e s on yeastmalt a g a r , oatmeal a g a r o r s a l t s - s t a r c h a g a r . One o b s e r v e r placed a e r i a l m a s s color in both the Gray and Red c o l o r - s e r i e s on these media, Good a e r i a l na ss growth is not found on glycerol-asparagine a g a r . Color tabs selected by o b s e r v e r s indicate a range f r o m light o r medium gray to yellowish o r brownish gray in the Gray s e r i e s , to grayish yellowish pink in the Red s e r i e s ; g r a y i s the predominant color. Reverse side of colony: Yellow to yellow-brown plus r e d (moderate orange to grayish r e d o r grayish reddish brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . The substrate mycelium pigment is not a pH indicator. Color in medium: Melanoid pigments a r e not produced in peptoneyeast-iron agar, tyrosine a g a r o r tryptone-yeast broth. Pigments a r e not found i n the medium i n yeast-malt a g a r , oatmeal a g a r o r s a l t s s t a r c h a g a r . A t r a c e of yellow pigment is found in glycerol-asparagine a g a r a f t e r 14-21 days; i t is not pH aensitive. D -Glucose, 5-arabinose, sucrose, _D-xylose, Carbon utilization: , i-inositol, _D-mannitol, _D-fructose, rhamnose and raffinose a r e a l l utilized f o r growth.
Streptomycee Rriseoruber Y amaguchi and Saburi. Description: Yamaguchi and Saburi 1955, 203 and 219-231. Not Streptomyces g r i s e o r u b e r (Ryabova and Preobrazhenskaya) P r i d h a m e t at. 1958, 69. Type strain: H-4650 E d . ) . ISP 5281 f r o m T. Yamaguchi as H-4650. ISP description by Group B-LO. Spore chain morphology: Section Spirales o r Retinaculiaperti. Open o r tight s p i r a l s a r e produced a t the ends of moderately long spore chains of LO to 50 o r m o r e s p o r e s p e r chain (Fig. 96a). Spirals a r e often incomplete, forming only 1 o r 2 t u r n s o r a hook at the end of the spore chain. Straight to flexuous chains without hooks o r s p i r a l s a r e a l s o p r e s e n t (Fig. 96b). This morphology is usually best on glycerol-asparagine a g a r , but may a l s o be observed on yeast-malt a g a r , oatmeal a g a r and e a l t s - s t a r c h a g a r . Spore surface: Smooth (Fig. 9 7 ) . Color of colony: Aerial m a s s color is i n the Red o r G r a y c o l o r - s e r i e s on yeast-malt agar, oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerolaeparagine a g a r . The color is represented by tab 5fe (light grayish reddish brown) f r o m the Gray c o l o r - s e r i e s and tab 5ge (also light grayi s h reddish brown) f r o m the Red color-serieB. Reverse aide of colony: Yellow to yellow-brown plus r e d (reddish orange o r reddieh brown on yeast-malt a g a r , oatmeal a g a r and s a l t s s t a r c h a g a r ; tight yellowish pink on glycerol-asparagine agar). Subs t r a t e mycelium pigment is a pH indicator changing f r o m reddish orange
Figure 94. A. griseoruber Ryabova and Preobrazhenskaya. Spiral spore chains @ 400) on yeast-malt agar, 14 days. Figure 95. A. griseoruber Ryabova and Preobrazhenskaya. Spiny spores; electronmicrograph from 16 day culture on yeast-malt agar. Figure 96. S. griseoruber Yamaguchi and Saburi. Spiral or R A spore chains on glycerol-asparagine a t a r , 14 days: (a) Open a n x i g h t spirals a t ends of long chains (X 580); (b) Straight t o glexmua chains and some terminal hooks o r spirals (X 230). l6 Figure 97. S. griseoruber Yamaguchi and Saburi. Smooth spores; electronmicrograph from 21 day culture on yeast-malt agar. l7
332
INTERNATIONAL
JOURNAL
to purple with addition of 0.05 N NaOH and f r o m reddish orange to yellowish orange with addition of 0.05 N HC1. Color i n medium: Melanoid pigments a r e produced in peptone-yeasti r o n a g a r , tyrosine a g a r and tryptone-yeast broth. Yellow to r e d pigment (depending upon pH) may be found i n the medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . This pigment is pH sensitive, changing f r o m orange to purple with addition of 0.05 N NaOH and f r o m orange t6 yellow with addition.of 0 . 0 5 N HC1. Carbon uti lization : _D -G luco s e , _L -a r a bin0 s e , ,D - xy lo s e ,i - i n o sito 1, D-fructose and rhamnose a r e utilized f o r growth. No growth o r only t r a c e of growth on s u c r o s e , _D-mannitol and raffinose. Streptomyces Kriseoviridis Anderson, Ehrlich, Sun and Burkholder Description: Streptomyces griseoviridus (91c) Anderson e t at. 1956, 109-115. (Spelling changed to 5 . g r i s e o v i r i d i s by Waksman 1961, 225, and by L. E. Anderson, personal communication, Sept., 1967. ) Type ISP 5229 f r o m L.E. strain: A 9071 = P-D 04955 = NRRL 2427 (%). Anderson a s P - D 04955 = NRRL 2427, IMRU 3735. ISP description by Group B -6. Spore chain morphology: Section S p i r a l e s (Fig. 98). Some open s p i r a l s have only one o r two t u r n s o r a r e poorly developed suggesting F U morphology. Spore chains moderately long with LO to 50 s p o r e s p e r chain. Spore surface: Smooth (Fig. 99). Color of colony: A e r i a l m a s s color in the Red c o l o r - s e r i e s (grayish yellowish pink to pale orange yellow) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . One o b s e r v e r placed The original description t h i s culture i n the Yellow c o l o r - s e r i e s . (Anderson G . , 1956) included the following statement: "Aerial mycelium i s white to pink-tan, pink-gray, o r brown o r occasionally greenish". The greenish color w a s not observed on ISP media. R e v e r s e side of colony: Grayed yellow to olive brown o r d a r k brown on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerola s p a r a g i n e a g a r . Substrate mycelium pigment i s not a pH indicator o r is changed only slightly by 0.05 N NaOH o r HCl. Color i n medium: Melanoid pigments a r e not found i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment i n medium, o r only t r a c e of yellow, in yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: D,-Gluco se, &-arabinose, _D-xylo s e, _D -mannito 1 and rhamnose a r e utilized f o r growth. Utilization of _D-fructose i s doubtful. No growth o r only t r a c e of growth on s u c r o s e , i - i n o s i t o l o r raffinos e.
Streptomyces g r i s e u s (Krainsky) Waksman and Henrici. Descriptions: Actinomyces g r i s e u s Krainsky 1914, 682; Actinomyces g r i s e u s (Krainsky) Wakeman 1916, 119 and 128; Waksman 1919, 123-125; Streptomyces g r i s e u s (Krainsky) Waksman and Henrici 1948, 948. (See a l s o s t reptomy cini K r a si l'nikov 1949, LOO ; Actinomyces globieporus va r . Xctinomyees etreptomycini Krasil'nikov 1957, 598). Type strain: IMRU 3463 tneotype, Waksman 1961, 226). ISP 5236 f r o m S.A. Waksman a s IMRU 3463. ISP description by Group B-LO. Spore chain morphology: Section Rectiflexibiles (Fig. LOO). Mature
Figure 9 8 . S. griseoviridis. Spiral spore chains (X 500) on oatmeal agar, 22 days. Is Figure 99. S . griseoviridis. Smooth spores, Au/Pd shadowed; electron micrograph from 21 day culture on oatmeal agar. yI spore chains (X 230) on yeast-malt agar, Figure 100. S. griseus. 14 days. Figure 101. 2. griseus. Smooth spores; electron micrograph from 14 day culture on yeast-malt agar. l6
334
s p o r e chains moderately long with LO to 50 s p o r e s p e r chain. This morphology is s e e n on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 101 ). Color of colony: A e r i a l m a s s color i n the Yellow c o l o r - s e r i e s (2db, pale yellow, o r 1 1/2 db - 1 1/2 ec, pale greenish yellow) on y e a s t m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine agar. R e v e r s e side of colony: No distinctive pigments (grayed yellow to olive brown o r light yellowish brown) on yeast-malt a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e f o r m e d i n tyrosine a g a r , but not i n peptone-yeast-iron a g a r o r tryptone-yeast broth. No pigments other than t r a c e s of yellow a r e found i n the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: Q-Glucose, _D-xylose, _D-mannitol a n d _D-fructose a r e utilized f o r growth. No growth o r only t r a c e of growth on &-arabinose, s u c r o s e , i-inositol, rhamnose o r raffinose. Streptomyces h e r b a r i c o l o r Kawato a n d Shinobu. Description: Kawato and Shinobu 1959, 114-119. Type strain: 608 (ibid.) ISP 5123 f r o m R. Shinobu as 608. ISP description by Group B-12. Spore chain morphology: Section Rectiflexibiles (Fig. 102). Mature s p o r e chains generally LO to 50 s p o r e s p e r chain on y e a s t - m a l t a g a r and glycerol-asparagine a g a r . Sporutating a e r i a l mycelium is poorly developed o r a b s e n t on oatmeal a g a r a n d s a l t s - s t a r c h a g a r . Spore surface: Smooth (Fig. 103). Color of colony: A e r i a l m a s s color i n the G r a y c o l o r - s e r i e s on glycerol-asparagine a g a r and y e a s t - m a t t a g a r . A e r i a l mycelium i s poorly developed on a l l ISP media; mass color cannot be determined on oatmeal a g a r or s a l t s - s t a r c h a g a r . R e v e r s e side of colony: Substrate mycelium may be grayish yellow to olive brown or brown o r i t may contain a n additional pH sensitive pigment with a color range f r o m pale purple o r p a l e purplish pink to g r e e n o r g r e e n i s h blue. When this pigment i s present, addition of 0.05 N NaOH changes color f r o m violet-red to g r e e n o r blue a n d this change is r e v e r s e d by addition of 0.05 N HC1. The pH sensitive mycelial pigment was found by one o b s e r v e r i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e not f o r m e d o r o c c u r only i n t r a c e amounts i n peptone-yeast-iron a g a r and tyrosine a g a r . Pigment m a y o r may not be f o r m e d i n the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . When formed, this pigment shows the s a m e pH range found i n the t r a n s i e n t s u b s t r a t e pigment. Carbon utilization: 2-Glucose, &-arabinose, s u c r o s e , _D-xylose, D-fructose and raffinose a r e utilized f o r growth. No growth o r only f growth on i-inositol, _D-rnannitol and rhamnose. trace o Streptomyces humidus Nakazawa, Shibata, Tanabe and Yamamoto. Descriptions: Nakazawa e t at. 1958, 321-324; a l s o Nakazawa e t at. 1960, U. S. P a t e n t s 2,931,756 and 2,950, 277. Name a n d antibiotic production only, without description: Nakazawa and Shibata & I m a m u r a -e t at. 1956, 648-653. Type strain: 23572 (Nakazawa e t a t , , H.).
F i g u r e 102. S . herbaricolor. epore chains (X 625) on yeast-malt agar, 14-day 8 . F i g u r e 103. S . herbaricolor. Smooth spores; electron micrograph f r o m 14 day c z t u r e on yeast-malt a g a r . 2Z F i g u r e 104. S . humidus. Spiral spore chains (X 600) on oatmeal agar, 14 days.76 F i g u r e 105. S. humidus. Smooth spores; electron micrograph f r o m 2 1 day c z h x a t m e a l a g a r . F i g u r e 106. S . indigoferus. Portion o f colony (X 400) showing R F spore chains a t e d g e of colony (top) and coremia on colony s u r f a c e
''
*'
F i g u r e 1 0 7 . 2. indigoferus. Smooth s p o r e s ; electron micrograph from 14 day culture on yeast-malt a g a r . 30
(bottom).
3 36
INTERNATIONAL
JOURNAL
ISP 5263 f r o m E. L e s s e l , A m e r . Type Cult. Cot. as ATCC 12760 f r o m Takeda Pharm. Ind., LTD. as 23572 = P O 3520. ISP description by Group B -10. Spore chain morphology: Section Spirales with many incomplete s p i r a l s resembling morphology (Fig. 104). Spore chains s h o r t to moderately long with LO to 50 s p o r e s p e r chain. T h i s morphology i s seen on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerola s p a r a g i n e a g a r , Spore surface: Smooth (Fig. 105). Special morphological c h a r a c t e r i s t i c s : Although the original d e s c r i p tion r e p o r t s production of black, moist (hygroscopic) a r e a s on m a t u r e a e r i a l mycelium, this was not r e c o r d e d on ISP r e p o r t s . Color of colony: A e r i a l m a s s color i n the G r a y o r Red colors e r i e s (3fe o r 31i, brownish gray, f r o m the G r a y c o l o r - s e r i e s to 5dc o r 5cb, g r a y i s h yellowish pink, i n the Red c o l o r - s e r i e s ) . R e v e r s e side of colony: No distinctive pigments (colorless, pale o r grayish yellow o r tight olive gray) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . Color i n medium: Me tanoid pigments not produced i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment, o r only t r a c e of yellow pigment, i s found i n the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: 9-Glucose, k a r a b i a o s e , _D-xylose, i-inositol, D-mannitol, _D-fructose and r h a m n o s e a r e utilized f o r growth. N o growth o r only t r a c e of growth on s u c r o s e and raffinose.
Streptomyces indigoferus Shinobu a n d Kawato. Descriptions: Shinobu and Kawato 1960b, 49-33. Type strain: 709 @bid.). ISP 5124 f r o m R. Shinobu a s 709. ISP description by Group B-5. Spore chain morphology: Section Rectiflexibiles. Mature s p o r e chains moderately long with LO to 5 0 , o r s o m e t i m e s m o r e than 50, s p o r e s p e r chain. A e r i a l mycelium is poorly developed on a l l ISP media and is obscured by extensive c o r e m i a formation on y e a s t - m a t t a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r (Fig. 106). Spore s u r face: Smooth (Fig. 107). Color of colony: A e r i a l mass color often cannot be determined because of inadequate a e r i a l growth on oatmeal a g a r o r because of extensive c o r e m i a formation on y e a s t - m a l t a g a r , s a l t s - s t a r c h a g a r aad glycerol-asparagine a g a r . When adequate a e r i a l mycelium i s formed, i t i s i n the Gray c o l o r - s e r i e s on yeast-malt a g a r and oatmeal a g a r . A e r i a l mass c o l o r i n the G r a y c o l o r - s e r i e s w a s a l s o found on glycerol- u r e a a g a r and soil e x t r a c t a g a r by one o b s e r v e r . R e v e r s e side of colony: Substrate mycelium i s usually nearly c o l o r less, grayish yellow o r d a r k brown on y e a s t - m a l t a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . A t r a n s i e n t blue to r e d pigment (depending on pH) may a l s o b e found i n the mycelium on t h e s e media. This pigment changes f r o m blue o r c o l o r l e s s to r e d with addition of 0.05 N HC1. Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r a n d tryptone-yeast broth, but a r e not f o r m e d i n tyrosine a g a r . P i g m e n t s o t h e r than melanoids w e r e not found i n the medium i n y e a s t m a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine agar. Carbon utilization: _D-Glucose a n d _L-arabinose a r e utilized for
SYSTEMATIC
BACTERIOLOGY
3 37
growth. Utilization of _D-xylose is doubtful. No growth o r only t r a c e of growth on sucrose, i-inositol, Q-mannitol, _D-fructose, rhamnose or raffinose. Actinomyces janthinus Artamonova and Krasillnikov. Description: Artamonova and Krasil'nikov 1960, 334 (see a l s o English translation, type s t r a i n selected by 3-7 and 300-306). Type strain: 117 N.A. Krasil'nikov, personal communication. ISP 5206 f r o m N-A. Krasillnikov a s INMI 117 = RIA 659. ISP description by Group B-8. Spore chain morphology: Section S p i r a l e s (Fig. LOB). Spore chains moderately long with LO to 50, or sometimes m o r e than 50, s p o r e s p e r chain. This morphology i s seen on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Spiny (Fig. 109). Color of colony: A e r i a l m a s s color i n the Red o r White c o l o r - s e r i e s on y e a s t - m a l t a g a r and s a l t s - s t a r c h a g a r ; White c o l o r - s e r i e s on oatmeal a g a r and White o r G r a y c o l o r - s e r i e s on glycerol-asparagine a g a r . The n e a r e s t color tabs i n the Red c o l o r - s e r i e s a r e 5dc, 5ca, 5cb and 6ec (all grayish yellowish pink). R e v e r s e side of colony: No distinctive pigments (colorless to grayed yellow or pale yellow) on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r or glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r and tryptone-yeast broth, but not in tyrosine a g a r . Yellow pigment is found i n the medium i n s a l t s - s t a r c h a g a r . This pigment i s not pH sensitive when tested with 0.05 N NaOH or HCI. Carbon utilization: 2-Glucose, &-arabinose, sucrose, _D-xylose, i-inositol, _D-mannitol, _D-fructose, rhamnose and raffinose a r e a l l utilized f o r growth.
(w);
Str eptoverticillium (SLC) ( B a r r and Carman) Baldacci, Farina and Locci. Descriptions: Streptomyces kentuchensis B a r r a n d Ca r m a n 1956, 2 86 2 89 ; St reptovertici 1lium kentuchense B a lda cci, Farina and Locci, 1966, 161. Type strain: 5x1 RCV a s Streptomyces kentuckensis B a r r and C a r m a n (ibid).ISP 5052 f r o m F . S . B a r r a s ATCC 12691 = 5x1 RCV. I S P description by Group B-1. Spore chain morphology: Umbellate Monoverticillate (= Streptomyces Section Verticillati, biverticillate) (Fig. 110). Mature spore chains moderately long with LO to 50 s p o r e s p e r chain. This morphology i s s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerola s p a r a g i n e a g a r . Spore surface: Smooth (Fig. 111). Color of colony: A e r i a l m a s s color i n the Red c o l o r - s e r i e s on y e a s t m a l t a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r ; Red or White colors e r i e s on glycerol-asparagine a g a r . R e v e r s e side of colony: No distinct pigments (grayed yellow on oatm e a l a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r to brown on y e a s t - m a tt a g a r ) . Color i n medium: Melanoid pigments a r e produced i n peptone-yeasti r o n a g a r and tryptone-yeast broth, but.not i n tyrosine a g a r . No pigment i s found i n the medium i n yeast-matt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r or glycerol-asparagine a g a r . Carbon utilization: _D-Glucose, L-inositol a n d fructose a r e utilized f o r growth; utilization of _D-xylose i s doubtful. No growth or only t r a c e
Figure 108. A . janthinus. Spiral spore chains (X 1000) on oatmeal agar, 14 days.n Figure 109. _A. janthinus. Spiny spores; electron micrograph from 14 day culture on oatmeal agar. l 9 Figure 110. Streptoverticillium kentuchense. E V (umbellatemonoverticillate) morphology (X 640) on yeast-malt agar, 14 days.' Figure 1 1 1 . Streritoverticilliumkentuchense. Smooth apores; electron micrograph from 21 day culture on oatmeal agar.
--
'
SYSTEMATIC of growth on L_-arabinose, raffinose.
BACTERIOLOGY _D-mannitol, rhamnose and
3 39
sucrose,
Actinomyces lavendocolor Kuchaeva, Krasil'nikov, Taptikova and Gesheva. Description: Kuchaeva et at. 1962, 103-124. Type strain: INA 4518 (ibid.). ISP 5216 f r o m N.A. Krasil'nikov a s INA 4518 = RIA 749. ISP description by Group B-12. Spore chain morphology: Section Retinaculiaperti o r E. Spore chains a r e long, often with m o r e than 50, s p o r e s p e r chain (Fig. 112). This morphology is seen on yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 113). Color of colony: A e r i a l m a s s color i n the Red c o l o r - s e r i e s on yeastm a l t a g a r , oatmeal a g a r a n d s a l t s - s t a r c h a g a r . A e r i a l mycelium is poorly developed on glycerol-asparagine a g a r . R e v e r s e side of colony: No distinct pigments (nearly c o l o r l e s s to pale yellow o r v e r y pale yellowish pink) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e f o r m e d in peptone-yeasti r o n a g a r and usually i n tyrosine a g a r a n d tryptone-yeast broth. P i g m e n t s other than melanoids a r e not found i n the medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-Glucose, &-arabinose, 2-xylose, a n d i inositol a r e utilized f o r growth. T r a c e of growth on sucrose, _D-mannitot, _D-fructose, rhamnose and raffinose r e s e m b l e s growth on the carbon f r e e control. Actinomyces lavendofoliae Kuchaeva, Krasillnikov, Taptikova and Gesheva. Description: Kuchaeva e t at. 1962, 103-124. Type strain: INA 3613 ( K d . ) . ISP 5217 f r o m N.A. Krasillnikov a s INA 3613 = RIA 750. ISP description by Group B-12. Spore chain morphology: Section Retinaculiaperti I Long, f Lexuous s p o r e chains often terminate i n s p i r a l s o r hooks and loops of wide d i a m e t e r (Fig. 114). Mature s p o r e chains have LO to 50, o r often m o r e than 50, s p o r e s p e r chain. This morphology i s seen on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 115). Color of colony: A e r i a l m a s s color i n the Red c o l o r - s e r i e s (Grayish yellowish pink o r yellowish pink) on yeast-malt a g a r , oatmeal a g a r and salts-starch agar. R e v e r s e side of colony: No distinctive pigment (pale yellow to light yellowish brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol asparagine a g a r . Color i n medium: Melanoid pigments are f o r m e d in peptone-yeasti r o n a g a r and usually i n tyrosine a g a r and tryptone-yeast broth. P i g m e n t s other than melanoids are not f o r m e d i n the medium in yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-Glucose, &-arabinose, _D-xylose and iinositol a r e utilized f o r growth. Utilization of f r u c t o s e is doubtful. No growth o r only trace of growth on s u c r o s e , e m a n n i t o l , rhamnose o r raffinose.
F i g u r e 112. A. lavendocolor. Very long flexuous chains of m o r e than 50 s p o r e s ( 5 4 8 5 ) on s a l t s - s t a r c h a g a r . 37 Loops and hooks, r e p r e sentative of F U cultures a r e a l s o reported for this s t r a i n . F i g u r e 113. 4. lavendocolor. Smooth s p o r e s ; electron micrograph f r o m 14 day culture on s a l t s - s t a r c h a g a r . " F i g u r e 114. A. lavendofoliae. R A s p o r e chains with t e r m i n a l s p i r a l s o r loops onGatmea1 a g a r , 21 days: (a) (X 300); (b) ( X 1200).23 F i g u r e 115. A. lavendofoliae. Smooth spores: electron micrograph from 14 day culture on s a l t s - s t a r c h a g a r . "
341
Actinomyces levoris Nikitina. Descriptions: Nikitina 1958, 142; Korenyako, Krasillnikov and Nikitina 1960, 116-L32 (see a l s o Engl. Transl., 111-127). Type strain: INMI 2725 (Ibld.; type s t r a i n selected by N.A. Krasil'nikov, personal communication). ISP 5202 f r o m N.A. Krasillnikov a s INMI 2725 = RIA 338. ISP description by Group B-5. Spore chain morphology: Section Rectiflexibiles. Mature s p o r e chains generally long with LO to 50, o r often m o r e than 50, s p o r e s p e r chain (Fig. 116). This morphology i s seen on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . Spore surface: SmoQth (Eig. 117). Color of colony: A e r i a l m a s s color i n Yellow c o l o r - s e r i e s on yeastm a l t a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Aerial mycelium may be inadequate f o r color observation on oatmeal a g a r . Reverse side of colony: No distinctive pigment (Grayish yellow o r grayish greenish yellow to orange-yellow o r yellowish brown) on yeastm a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine agar. Color in medium: Melanoid pigments a r e not f o r m e d i n peptoneyeast-iron a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment i s found i n the medium i n yeast-malt agar, oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon uti liza tion: ,D -G luco se , _L-a rabino se, ,D -xyLo se, _D -mannit01 and 2 - f r u c t o s e a r e utilized for growth. No growth o r only t r a c e of growth on sucrose, i--inositol, rhamnose and raffinose. Streptomyces lilacinus Nakazawa, Tanabe, Shibata, Miyake a n d Takewaka. Descriptions: Nakazawa e t at. 1956, 81; Nakazawa e t at. 1957, 111-115. Type strain: 2305 u ) F I S P 5254 f r o m K. Nakazawa a s IF0 3944 = 2305. ISP description by Group B-LO. Spore chain morphology: Section VerticiLlati, umbellate monoverticillate (biverticillate), (Fig. 118). Spore chains a r e short, usually with 3-10 s p o r e s p e r chain. This morphology is seen on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 119). Color of colony: A e r i a l m a s s color i n the Red c o l o r - s e r i e s ( grayish yellowish pink) on yeast-malt a g a r , oatmeal a g a r , salts-starch a g a r and g lyc e r o 1-a spa r a g ine a g a r R e v e r s e side of colony: Yellow-brown plus r e d (grayish reddish brown to strong brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ; this pigment is somewhat pH sensitive changing f r o m brown to reddish brown with addition of 0 . 0 5 N NaOH and f r o m brown to yellowish brown with addition of 0.05 N H C 1 . Color in medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r a n d tryptone-yeast broth. Red pigment is found i n the medium i n yeast-malt agar, oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . This pigment is pH sensitive when tested with 0.05 N NaOH o r HCl, showing the s a m e change noted f o r s u b s t r a t e color. Carbon utilization: 2-Glucose i s utilized f o r growth; r e p o r t s v a r y on positive utilization ofi-inositol. No growth o r only t r a c e of growth on _L-arabinose, sucrose, g-xylose, _D-mann.itol, _D-fructose, rhamnose a n d raffinose.
342
Actinomyce s longisporus K r a sil'nikov. Description: K r a s i l'nikov 1941, 47. S t r e tom ces !ongisporus (Krasil'nikov) Waksman 1953, 39. Type s t r 4417 a 56 i (neotype, w Preobrashenskaya, p e r s o n a l communication: N.A. Krasil'nikov, p e r s o n a l communication, January, 1968). ISP 5166 f r o m T.P. P reobrazhenskaya a s 4417/56. ISP description by Group B-1. Spore chain morphology: Section S p i r a l e s (Fig. 120). Mature s p o r e chains moderately short, usually LO to 50 s p o r e s p e r chain. Spore surface: Spiny (Fig. 121). (One o b s e r v e r found smooth s p o r e s . ) Color of colony: A e r i a l m a s s color i n the White c o l o r - s e r i e s on y e a s t - m a t t a g a r , oatmeal a g a r and glycerol-asparagine a g a r ; White o r Red (5cb g r a y i s h yellowish pink) c o l o r - s e r i e s on s a l t s - s t a r c h a g a r . R e v e r s e side of colony: No distinctive pigments (light o r grayish yellow on oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r to olive brown o r brown on y e a s t - m a l t agar). Color i n medium: Melanoid pigments a r e produced i n peptone-yeast i r o n a g a r , tyrosine a g a r a n d tryptone-yeast broth. P i g m e n t s other than melanoids a r e not found i n the medium i n yeast-malt a g a r , oatm e a l a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon uti liza tion: _D-G lucose, k-ara binose, sucrose, E-xy lo se, i - inositol, 2-mannitol, _D-fructose, r h a m n o s e and raffinose a r e a l l utilized for growth. Actinomyce s longi s p o r u s -f lavus K r a sil'nikov. Descriptions : Krasil'nikov 1941, 30. Type strain: 8 1/53 (neotype, Preobrazhenskaya, pe r sona 1 communication ; K r a si 1' nikov, p e r s ona 1 co mm uni cati on). Streptomyces longisporoflavus Waksman 1953, 94; Streptomyces Longisporus flavus (Krasil'nikov) P r i d h a m e t at. 1958, 66. ISP 5165 f r o m T. P. Preobrazhenskaya a s 5165. ISP description by Group A-9. Spore chain morphology: Section S p i r a l e s Open s p i r a l s ; s p o r e chains resembling morphology a r e a l s o common (Fig. 122). Mature s p o r e chains a r e long with LO t o 50 s p o r e s p e r chain; longer chains a r e often observed. This morphology may be observed on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 123). Color of colony: A e r i a l n a s s color i s i n Yellow c o l o r - s e r i e s on s a l t s - s t a r c h a g a r . A e r i a l mycelium is poorly developed o r absent on other ISP media. R e v e r s e side of colony: No distinct pigments (colorless, grayish yellow o r light g r a y i s h yellowish brown) on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e not f o r m e d i n peptoney e a s t - i r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment found i n medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-Clucose, L.-arabinose, _D-xylose, D - f r u c t o s e and r h a m n o e a r e utilized f o r growth. No growth o r only t r a c e of growth on s u c r o s e and raffinose. Utilization of i - i n o s i t o l and 2mannitol i s doubtful.
St reptomyce s luteoverticit latus Shinobu. Description: Shinobu 1956, 84-93. Type strain: 486 (xd.). ISP 5038 f r o m R. Shinobu as 486. ISP description by Group A-8.
Figure 116; A. levoris. RF spore chains (X 300) on glycerol-aeparagine agar, 5 Z y s i Figure 117. _A. levoris. Smooth spores; electron micrograph from 32 day culture on salts- starch agar. Figure 118. 2. lilacinus. BIV (umbellate monoverticllate) morphology (X 400) on salts-starch agar, 14 days. Figure 119. 2. lilacinus. Smooth spores; electron micrograph from 21 day culture on oatmeal agar. Figure 120. _A. lonfisporus. Spiral spore chains (X 640) on oatmeal agar, 21 days. Figure 121. ,A. longisporus. Spiny spores; electron micrograph from 14 day culture on salts-starch agar.
344
S p o r e chain morphology: Section Verticillati, umbellate monoverticillate (biverticillate, F i g . 124). Mature s p o r e chains generally 10 t o 20 o r m o r e s p o r e s p e r chain a f t e r 21 days on s a l t s - s t a r c h a g a r or y e a s t matt a g a r . A e r i a l mycelium is poorly developed or a b s e n t on y e a s t matt a g a r , oatmeat a g a r a n d glycerol-asparagine a g a r . Absence of good a e r i a l mycelium on v a r i o u s media was a l s o noted by Shinobu i n h i s original description &, c&.). Spore surface: Smooth. Color of colony: Mature a e r i a l mycelium i n the Red o r Yellow colors e r i e s on s a l t s - s t a r c h a g a r ; i t is absent o r poorly developed (White o r Yellow) on other ISP media. R e v e r s e side of colony: No distinctive pigments (light brown to grayed yellow) on y e a s t - m a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r g Lyce r o 1-a sparagine a g a r . Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r and tryptone-yeast broth. Yellow pigment i s found in medium i n y e a s t - m a l t a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r a f t e r L4-21 days; it i s not a pH indicator. Carbon utilization: _D-Glucose, i-inositol a d D - f m t t o s e a r e utilized f o r growth. No growth o r only t r a c e of growth on &-arabinose, s u c r o s e , _D-xylose, _D-mannitol, rhamnose and raffinose. Streptomyces m a s h u e n s i s Sawazaki, Susuki, Nakamura, Kawasaki, Y a m a s h i t a , Isono, Anzai, Serizawa a n d Sekiyama. Description: Sawazaki et at. 1955, 44-45. Type strain: 449 ( S . Suski, p e r s o n a l communication). ISP 5221 f r o m S. Susuki a s 449. ISP description by S r o u p B-5. Spore chain morphology. Section Verticillati, umbellate monoverticillate (biverticillate, Fig. 125). Spore chains a r e s h o r t with 3 to LO, o r s o m e t i m e s m o r e than LO, s p o r e s p e r chain. Spore surface: Smooth (Fig. 126). Color of colony: A e r i a l mass color i n the Red c o l o r - s e r i e s on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d g l y c e r o l - a s p a r agine a g a r . Mature aerial mycelium m a y be poorly developed on glycerol-asparagine a g a r . R e v e r s e side of colony: No distinctive pigments (grayish yellow to orange yellow on o a t m e a l a g a r , s a l t s - s t a r c h a g a r and glycerolasparagine a g a r ; olive brown to brown on y e a s t - m a l t a g a r ) . Color i n medium: Melanoid pigments a r e produced i n peptone-yeasti r o n a g a r a n d tryptone-yeast broth, but not i n t y r o s i n e a g a r . No pigment i s found i n the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-Glucose, s u c r o s e and _D-fructose a r e utilized for growth. Utilization of 1-inositol i s doubtful. No growth o r only t r a c e of growth on &-arabinose, _D-xylose, _D-mannitol, r h a m n o s e and raffinose. Streptomyces m a t e n s i s Margalith, B e r e t t a and Timbal. Description: Margalith et at. 1959, 71-75. Type s t r a i n : ME/17 k d , ) . ISP 5188 f r o m L.G. Silvestri, Lepetit S.p.A. as Lep. M-ME/l7. ISP d e s c r i p tion by Group A-13. Spore chain morphology: Section Spirales. Open s p i r a l s a r e the dominant f o r m s (Fig. 127); s p o r e chains suggesting primitive s p i r a l s of cultures o r flexuous chains of = a r e a l s o present. Mature s p o r e chains moderately long with LO to 50 o r m o r e s p o r e s p e r chain.
Figure 122. A. longisporus-flaws. (a) Spiral spore chains (X 400) on o a t m e a l a g a r , 14 days. (b) Terminal spirals and hooks on yeastmalt agar, 14 days. Figure 123. A. longisporus-flaws. Smooth spores; electron micrograph from 21 day culture on salts-starch agar. Figure 124. S. luteoverticillatus. BIV (umbellate monoverticillate) morphology on yea st -malt agar? Figure 125. S. mashuensis. BIV (umbellate monoverticillate) morphology (X 400) oatmeal agar, 14 days. Figure 126. S. mashuensis. Smooth spores; electron micrograph from 10 day c z t u r e on yeast-malt agar.'
''
''
'
3 46
This morphology is seen on yeast-malt agar, oatmeal agar, saltsstarch a g a r and glycerol-asparagine agar. Spore surface: Spiny. Spines a r e short and inconspicuous (Fig. 128); some smooth spores may be seen. Color of colony: Mature a e r i a l m a s s color in the Gray color-series (color tab 3ge, tight grayish yellowish brown; to 5fe, light grayish reddish brown) on salts-starch a g a r ; Gray o r White color-series on yeast-malt a g a r and glycerol-asparagine agar. One observer reports tab 5dc (grayish yellowish pink) f r o m the Red color-series for growth on yeast-malt agar, oatmeal a g a r and salts-starch a g a r . This trace of r e d (or violet) pigment in a e r i a l color is also noted in the original description (Margalith et at. 1959). Reverse side of colony: Red pigments in mycelium modify charact e r i s t i c substrate colors to d a r k reddish brown on yeast-malt a g a r and salts-starch agar, tight grayish brown o r yellowish pink on oatmeal a g a r and glycerol-asparagine agar. The substrate mycelium pigment i s not a strong pH indicator. (One observer reports change from violet to pink with addition of 0.05 N HC1. ) Color in medium: Melanoid pigments a r e not found in peptone-yeastiron agar, tyrosine a g a r o r tryptone-yeast broth. Red o r violet pigments a r e found in yeast-malt agar, oatmeal a g a r and glycerol-asparagine a g a r . One observer reports a change from violet to red by addition of 0.05 N H C l . Carbon utilization: 2-Glucose, &-arabinose, _D-xylose, i-inositol, D-mannitol, _D-fructose and rhamnose a r e utilized for growth. N o growth o r only t r a c e of growth on raffinose. Utilization of sucrose i s doubtful.
Streptomycea mediocidicus Okami, Uta ha ra , Nakamura and Umezawa. Description: Okami et at. 1954, 98-100. Type strain: 535-A1 (ibid.). H J 266. ISP description by ISP 5021 from Y. Okami a s 535-A1 = M Group B -2. Spore chain morphology: Sporulating a e r i a l mycelium is not produced on any ISP media o r on 15 additional media tested by cooperators. Single conidia were observed on the substrate mycelium on glucoseasparagine a g a r and Waksman's s t a r c h a g a r B (Fig. 129). Masses of c r y s t a l s formed on the substrate growth on yeast-malt a g a r (Fig. 130) The original description (Okami et at. 1954) indicates that a e r i a l mycelium w a s only poorly developed o r absent on most media. Spore surface: not observed. Color of colony: No sporulating a e r i a l mycelium w a s found on any of the media tried by the cooperators. Reverse side of colony: Yellow to greenish yellow on yeast-malt agar, oatmeal agar, salts-starch a g a r and glycerol-asparagine agar. Thie pigment is not pH sensitive. Color in medium: Melanoid pigments a r e formed in peptone-yeastiron a g a r and tryptone-yeast broth, but may not be formed in tyrosine agar. Trace of yellow pigment may be found in the medium in yeastmalt agar, oatmeal agar, s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ; this pigment is not pH sensitive. Carbon utilization: D_-Glucose, &-inositol and _D-fructose a r e utilized f o r growth. No growth o r only t r a c e of growth on &-arabinose, sucrose, D-xylose, _D-mannitol, rhamnose and raffinose.
F i g u r e 127. -S. matensis. Spiral spore chains on yeast-malt agar. F i g u r e 128. 2. matensis. Spiny spores; spines a r e short and sparse. Electron micrograph f r o m 13 day culture on Gauze medium No. 1. F i g u r e 129. S. mediocidicus. Single s p o r e s on substrate mycelium: (a) On Waksmanls s t a r c h a g a r B, 21 days.te (b) On glucoeeasparagine agar, 10 days (X 1000). * F i g u r e 130. S. mediocidicus. Crystalline m a m a on surface of eubstratal growth (X 380) on yeast-malt agar, 21 days.
3 48
I N T E R N A T I O N A L J O U R NA L
Streptomyces melanogenes Sugawara and Onuma. Description: Sugawara and Onuma 1957, 138-142. Type strain: KITA V-1179 (T. Hata, personal communication). ISP 5192 f r o m Y. Yagisawa f r o m T. Hata, Kitasato Inst. a s KITA V-1179. ISP description by Group B-2. Spore chain morphology: Section Rectiflexibiles (Fig. 131). Mature s p o r e chains long with LO to 50, o r often m o r e than 50, s p o r e s p e r chain. This morphology is s e e n on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 132). Color of colony: A e r i a l m a s s color i n the Red c o l o r - s e r i e s (5cb, g r a y i s h yellowish pink) on oatmeal a g a r and s a l t s - s t a r c h a g a r ; White o r Gray c o l o r - s e r i e s on yeast-malt a g a r and s a l t s - s t a r c h a g a r . Mature sporulating a e r i a l mycelium may be poorly developed on y e a s t matt a g a r and s a l t s - s t a r c h a g a r . R e v e r s e side of colony: Grayed yellow to yellow-brown is usually modified by r e d (becoming d a r k brown to reddish brown) on yeast-malt a g a r , oatmeat a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ; t h i s pigment is not a pH indicator. Color i n medium: Melanoid pigments a r e f o r m e d in peptone-yeasti r o n a g a r , tyrosine a g a r and tryptone-yeast broth. Yellow o r r e d pigment is found in the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r . This pigment i s not pH sensitive when t e s t e d with 0.05 N NaOH o r HC1. Carbon utilization: _D-Glucose, &-arabinose, D-xylose, i-inositol, D-mannitol, 2 - f r u c t o s e and raffinose a r e utilized f o r growth. Reports v a r y on utilization of sucrose. No growth o r only t r a c e of growth with rhamnose.
Streptomyces misakiensis Nakamura. Description: Nakamura 1961, 86-89; name and antibiotic production, without description, i n Isono, Anzai and Suzuki 1958, 264-267. Type strain: 1755 (Nakamura, ISP 5222 f r o m S . Suouki, Inst. Phys. and Chem. Res., Tokyo a s 7617 = 1755. ISP description by Group B-6. Spore chain morphology: Section Rectiflexibiles (Fig. 133). Mature s p o r e chains generally tong with LO to 50, o r s o m e t i m e s m o r e than 50, s p o r e s p e r chain. This morphology i s s e e n on y e a s t - m a l t a g a r , oatm e a l a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore s u r f a u z . Smooth (Fig. 134). Color of colony: A e r i a l m a s s color i n the Gray c o l o r - s e r i e s on s a l t s - s t a r c h a g a r and glycerol-aeparagine a g a r ; G r a y o r Red colors e r i e s on y e a s t - m a l t a g a r and oatmeal a g a r . The m o s t representative color tab f r o m the Gray color-wheel i s 2dc (yellowish g r a y ) f o r a l l ISP media. One o b s e r v e r selected tab 5dc (grayish yellowish pink) f r o m the Red color-wheel a s the m o s t representative color on y e a s t malt a g a r and oatmeal a g a r . R e v e r s e side of colony: Yellow to yellow-brown i s modified by red, becoming brown to strong brown on yeast-malt a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r and orange to tight brown on oatmeal a g a r ; t h i s pigment is not a pH indicator. Color i n medium: Melanoid pigments a r e not formed in peptoneyeast-iron a g a r , tyrosine a g a r o r tryptone-yeast broth. Red pigment is found i n the medium i n yeast-matt a g a r , oatmeal a g a r and s a l t s s t a r c h a g a r . This pigment is not pH sensitive when tested with 0.05 NaOH o r HCl.
s.).
Figure 131. S. melanogenes. R F spore chains. (a) (X 380), Waksman's starch agar B, 2 1 days.'' (b)(X 500), oatmeal agar, 7 days.' Figure 132. S. melanogenes. Smooth spores; electron micrograph from 21 day culture on oatmeal agar. 2 9 Figure 133. S. misakiensis. R F spore chains (X 350) onyeadt-malt agar, 2 1-day s. Figure 134. S. misakiensis. Smooth spores; electron micrograph (Au/Pd axadowed) from 21 day culture on oatmeal agar.
350
INTERNATIONAL
JOURNAL
Carbon utilization: _D-GLucose, s u c r o s e , _D-fructose and raffinose a r e utilized f o r growth. N o growth o r only trace of growth with Aarabinose, _D-xylose, i-inositol, _D-mannitol and rhamnose. In the original description (Nakamura, K. ) L_-arabinose and _D-mannitot w e r e a l s o reported a s supporting growth with Czapek's a g a r a s b a s a l medium.
& .
Streptomyces naganishii Yamaguchi and Saburi. Description: Yamaguchi a n d Saburi 1955, 216-219 and 226. Type strain: H-4871 ISP 5282 f r o m T. Yamaguchi a s H-4871. ISP description by kd.). Group B -11. Spore chain morphology: Section S p i r a l e s (Fig. 135a, b). Mature s p o r e chains s h o r t to moderately long, usually with m o r e than LO s p o r e s p e r chain. This morphology is best developed on yeast-malt a g a r . Short s p o r e chains on s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r may f o r m incomplete s p i r a l s o r hooks (Fig. 135c). Sporulation on oatmeal a g a r is especially poor. A e r i a l hyphae m a y t e r m i n a t e i n atypical club-Like swellings instead of s p o r e s (Fig. 136). Spore surface: Smooth (Fig. 137). Color o f colony: A e r i a l m a s s color i n the G r a y o r Red c o l o r - s e r i e s on yeast-matt a g a r a n d on carbon utilization medium plus _D-Glucose. The sporulated a e r i a l growth on oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r is usually white. R e v e r s e side of colony: P a l e o r grayish yellow t o yellow-brown modified i n spots o r a t m a r g i n s by r e d (dark pink, purplish pink o r g r a y i s h reddish brown). S u b s t r a t e pigment is not a pH indicator. Color in medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r and tryptone-yeast broth i n two days, but a r e produced slowly o r not a t a l l i n tyrosine a g a r . A t r a n s i e n t r e d pigment i s found i n the medium in oatmeal a g a r a n d s a l t s - s t a r c h a g a r . This pigment is not pH sensitive when t e s t e d w i t h 0.05 N NaOH or HC1. Carbon utilization: _D-GLucose, _L-arabinose, _D-xylose, i-inositol, D-mannitol, _D-fructose and rhamnose a re utilized f o r growth. Utilization of raffinose i s doubtful. No growth o r only trace of growth with sucrose.
Streptoverticillium n e t r o p s i s (Finlay, Hochstein, Sobin a n d Murphy) Baldacci, Farina a n d Locci. Description: Streptomyces n e t r o p s i s , U. S. P a t e n t 2,586,762, Feb. 19,1952; name and antibiotic production, without description, i n Finlay, Hochstein, Sobin a n d Murphy 1951, 341-343. Streptoverticillium netropsis, Baldacci, F a r i n a a n d L o c c i 1966, 161-162. Type strain: Isolate 2937-6 = P f i z e r F . D . 4779 = NRRL 2268. ISP 5259 f r o m J. Routien, Chas. P f i z e r a n d Go., Inc. a s F. D. 4779 = NRRL 2268. ISP description by Group B-6. Spore chain morphology: Umbellate monoverticillate (= Streptomyces Section Verticillati, biverticillate). Whorls may be composed of s t r a i g h t (BIV), flexuous, o r s p i r a l (BBV-2) s p o r e chains (Fig. 138a, b, c). Mature s p o r e chains generally LO to 20 s p o r e s p e r chain. This morphology i s s e e n on y e a s t - m a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerolasparagine a g a r . Spore surface: Smooth (Fig. 139). Color of colony: A e r i a l m a s s color i n the Red c o l o r - s e r i e s on y e a s t m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine agar.
Figure 135. 2. na anishii. a) S iral spore chains ( X 200) on glycerolasparagine ag:r, 14 day(s. *Spiral chains (X 400) on yeast-malt agar. ' 9 ( c ) Short chains with incomplete spirals and hooks, yeastmalt agar, 14 days.36 Figure 136. S. na anishii Aerial hyphae terminating in club-like swellingc(%*ieast-malt agar. 36 Figure 137. 2. naganishii. Smooth spores; electron micrograph from 14 day culture on glycerol-asparagine agar.
Figure 138. Streptoverticillium netropsis. (a) E V (umbellate monoverticillate) morphology with straight chains forming the umbels ( X 500) on oatmeal agar, 24 days. Is (b) BIV with flexuous spore chains, ( X 500) on salts-starch agar, 2 4 x y s . l5 (c) BIV-S ( X 350); spore chains terminate in coils of one to two turns; salts-starch agar, 20 days.'' Figure 139. Stre overticillium netro sis. Smooth spores; electron m i c r o g r a e d ) f r z m 21 day culture on oatmeal agar.u
SYSTEMATIC
BACTERIOLOGY
353
R e v e r s e side of colony: No distinctive pigments (grayish yellow to olive brown o r brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerbl-asparagine a g a r . Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r , but not i n tyrosine a g a r . No pigment o r only a t r a c e of yellow o r brown pigment, is found i n the medium i n yeast-malt a g a r , oatmeal a g a r a n d s a l t s - s t a r c h a g a r . Pigment, if present, m a y be slightly pH sensitive changing f r o m pate yellow to pale pink when tested with 0.05 N NaOH. Carbon utilization: _D-Glucose, L-inositol and _D-fructose a r e utilized f o r growth. No growth o r only t r a c e of growth w i t h L_-arabinose, s u c r o s e , _D-xylose, _D-mannitol, rhamnose and raffinose. Actinomyces n i g r e s c e n s Sveshnikova. Description: Sveshnikova in Gauze e t at. 1957, 146-148. Type strain: 1800/54 ( E d . ; selected by G.F. Gauze and regarded by him a s identical with Streptomyces hygroscopicus, personal communication, Sept., 1964). Streptomyces nigrescene (Sveshnikova) P r i d h a m e t a t . 1958, 70. ISP 5276 f r o m G.F. Gauze a s 1800/54. ISP description by Group B -6. Spore chain morphology: Section Spirales (Fig. 140). Mature s p o r e chains moderately long with LO to 50, o r s o m e t i m e s m o r e than 50, s p o r e s p e r chain. This morphology i s s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth to warty (Fig. 141). Color of colony: A e r i a l m a s s color i n the G r a y c o l o r - s e r i e s on y e a s t m a l t a g a r , oatmeal a g a r , s a t t s - s t a r c h a g a r and glycerol-asparagine a g a r . In the original description ( S v e s h n i k o v a 9 .&.) emphasis i s placed on a n autolytic change of g r a y a e r i a l mycellium to a black shiny m a s s i n aging cultures. This phenomenon was not observed by ISP cooperators. R e v e r s e side of colony: No distinctive pigments (colorless to g r a y i s h yellow) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerolasparagine agar. Color i n medium: Melanoid pigments a r e not f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment i s found i n the medium i n yeast-malt a g a r , oatmeal a g a r , s a t t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: ~-GLUCOSe, sucrose, _D-xylose, i-inositol, D-mannitol, _D-fructose and raffinose a r e utilized f o r growth. No growth or only t r a c e of growth with -arabinose and rhamnose. Streptomyces nodosus Trejo. Description: T r e j o in Waksman 1961, 250; T r e j o and Bennett 1963, 436-439. Type strain: SC 2388 = ATCC .~ 14899 (Trejo, personal communication). ISP 5109 f r o m W. H. T r e j o a s SC 2388. ISP description by Group A-12. Spore chain morphology: Section Spirales. S p i r a l s on short spore chains may be poorly developed showing only a few t u r n s o r hooks of small d i a m e t e r ; o r tightly knotted s p i r a l s may be f o r m e d (Fig. 142). Spore chains a r e short, often with only 3 to LO s p o r e s p e r chain. This morphology i s seen on yeast-malt a g a r , s a l t s - s t a r c h a g a r and glycerola s p a r a g i n e a g a r . Spore surface: Smooth, but some wrinkles o r folds a r e found on the s p o r e surface (Fig. 143).
Figure 140. ,A. ni rescens agar, 21 days.
-%------
Figure 141. A. nigrescsns. Smooth spores (Au/Pd shadowed) with small suFface irregularities suggesting warts. Figure 142. 2. nodosus. Spirals on shqrt chains, some forming tight "knots" (X 400); salts-starch agar, 21 days. Figure 143. 2. nodosus. Smooth spores; electron micrograph from 14 day culture on salts-starch agar.
S i r a l spore chains (X 300) on salts-starch
''
*'
SYSTEMATIC
BACTERIOLOGY
355
Color of colony: A e r i a l m a s s color in the G r a y c o l o r - s e r i e s on yeast-malt a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r , R e v e r s e side of colony: Grayed yellow o r grayed greenish yellow to d a r k olive brown o r n e a r black on yeast-malt a g a r , oatmeal a g a r , sa lt s - s t a r c h a g a r and g lyce r o 1-a sparagine a g a r ; sub s t r a t e myce lium pigment i s not a pH indicator. Color i n medium: Melanoid pigments not f o r m e d in peptone-yeasti r o n a g a r , tyrosine a g a r and tryptone-yeast e x t r a c t broth. T r a c e s of r e d or yellow orange pigment may be found in the medium i n yeast-malt a g a r , oatmeal a g a r and glycerol-asparagine a g a r . One o b s e r v e r r e p o r t s this pigmert is pH sensitive changing f r o m r e d to g r e e n by addition of 0.05 N NaOH and f r o m g r e e n to blue-red by 0.05 N HC1. D - x y l o se, i - i n o sito 1, _D -mannito 1, Carbon uti t i za tion: _D -G Lucose, , D-fructose and rhamnose a r e utilized f o r growth. No growth or only trace of growth on s u c r o s e and raffinose. Utilization of .L-arabinoae is doubtful
(Arai, Nakada and Suzuki) BaldaCCi, S t r eptove r tici 1lium Farina and Locci. Descriptions: Streptomyces olivoreticuli A r a i e t a l . 1957, 435-441; Streptoverticillium olivoreticuli (Arai e t al. ) Baldacci, Farina and Locci 1966, 162. Type strain: LOO (ibid.) = I F M 1018, A r a i , p e r s o n a l communication. ISP 5105 f r o m T. A r a i a s I F M 1018 = LOO. ISP description by Group B-1. Spore chain morphology: Umbellate monoverticillate ( = Streptomyces Section Verticillati, biverticillate, Fig. 144). Mature spore chains generally s h o r t with 3 to LO s p o r e s p e r chain. This morphology m a y be observed on P r i d h a m and Gottlieb carbon utilization medium with glucose o r inositol, but good sporulation is usually not seen on yeastm a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 145). Color of colony: A e r i a l m a s s color i n the White c o l o r - s e r i e s on media supporting sporulating a e r i a l growth. In the absence of sporulation, the surface is pate yellow on yeast-matt a g a r , oatmeal a g a r and glycerol-asparagine a g a r and white on s a l t s - s t a r c h a g a r . R e v e r s e side of colony: No distinctive pigments (colorless to g r a y i s h yellowish brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r and tryptone-yeast broth, but not i n tyrosine a g a r . No pigment i s found i n the medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: Carbon utilization a p p e a r s to be unusually variable. Reports f r o m t h r e e o b s e r v e r s differ on utilization of each carbon s o u r c e tested and r e s u l t s v a r y on duplicate dishes i n the s a m e laboratory. The original description (Arai, e t a t . 1957) d i r e c t s attention to fluctuation of carbon utilization by ultra-violet i r r a d i a t e d s t r a i n s . This fluctuation a p p e a r s a l s o to be a c h a r a c t e r i s t i c of the type strain. Streptomyces olivoviridis Kuchaeva, Krasillnikov, Skryabin and Taptykova. Description: Kuchaeva e t at. 1960, 251 (see a l s o Engl. k a n s l . , 222-251). Type strain: INMI 1475 = RIA 661 ( E d . ; type s t r a i n ?elected by N . A . Krasil'nikov, personal communication). ISP 5211 f r o m N.A. Krasil'nikov a s INMI 1475 = RIA 661. ISP description by Group B-8.
Figure 144. Streptoverticillium olivoreticuli. BIV (umbellate monoverticillate) morphology (X 640) on P r i d h a m a n d Gottlieb carbon utilization medium plus inositol, 14 days. Figure 145. Streptoverticillium olivoreticuli. Smooth spores; electron micrograph from 28 day culture on salts-starch agar. a ) R F spore chains (X 300) on yeast-malt Figure 146. S. olivoviridis. ' agar, 7 days. (b) Flexuousand wavy chains with some tendency toward hooks and spirals; 14 day culture on oatmeal agar. l 3 Figure 147. 2. olivoviridis. Spiny spores; electron micrograph from 14 day culture on oatmeal agar. l 9
''
SYSTEMATIC
BACTERIOLOGY
357
Spore chain morphology: Section Rectiflexibiles (Fig. 146a). The predominantly flexuous o r wavy s p o r e chains m a y a l s o f o r m occasional hooks and open s p i r a l s ( r a r e ) resembling o r ,S morphology (Fig. 146b). Mature s p o r e chains generally long with LO to 50 s p o r e s p e r chain. This morphology i s seen on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Spiny (Fig. 147). Color of colony: A e r i a l m a s s color i n the Gray c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r ; a e r i a l mycelium i s poorly developed o r white on glycerol-asparagine a g a r . Reverse side of colony: Yellow to yellow brown is usually modified by green (olive brown to yellow g r e e n o r grayish green) on yeast-malt a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r ; no distinctive substrate mycelium pigment on glycerol-asparagine a g a r . Substrate mycelium pigment i s not a pH indicator. Color in medium: Melanoid pigments a r e not f o r m e d in peptoneyeast-iron a g a r o r tyrosine a g a r . Some coloration m a y appear i n medium i n tryptone-yeast broth. Yellow o r g r e e n pigment is found i n the medium in yeast-malt a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r . This pigment is not pH sensitive when tested with 0.05 N NaOH o r HC1. Carbon uti1ization:g- Glucose, 4 - a r a b i n o s e , _D-xylose, _D-maMitot, D-fructose and rhamnose a r e utilized f o r growth. No growth o r only t r a c e of growth with sucrose, I-inositol o r raffinose. Streptomyces phaeochromogenes (Conn) Waksman. Descriptions: Actinomyces pheochromogenus (sic ) Conn 1917a, 3, 16 (also reprinted i n Conn 1917b, 157,170); Streptomyces phaeochromogenus (Conn) Waksman and Henrici 1948, 943; Streptomyces phaeochromogenes (Conn) Waksman 1957, 778. Type strain: IMRU 3338 (Waksman, 1961). ISP 5073 f r o m S.A. Waksman a s IMRU 3338. ISP description by Group B-4. Spore chain morphology: Section Rectiflexibiles (Fig. 148). Mature s p o r e chains generally straight chains of LO to 50, o r sometimes m o r e than 50, s p o r e s p e r chain. This morphology is seen on yeast-malt a g a r , oatmeal agar, s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . S p i r a l s were reported in Wakeman's 1919 description, but not i n Conn's original description (1917, 3 . &t. ) Subsequent observations reported i n Waksman, 1961, note frequent occurrence of straight spore chains and absence of spirals. Spore surface: Smooth (Fig. 149). Color of colony: A e r i a l m a s s color usually i n the Red c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerolasparagine a g a r . Most representative color tabs f r o m the Red colors e r i e s w e r e 3ca (pale orange yellow) and 5ca o r 5cb (light yellowish pink to grayish yellowish pink). A e r i a l na ss color may a l s o b e white o r pate yetlow on oatmeal a g a r and pate yellow on s a l t s - s t a r c h a g a r a n d g Lyc e r o 1-a spa ragine a g a r . Reverse side of colony: No distinctive pigments (pale yellow o r grayish yellow on oatmeal a g a r and s a l t s - s t a r c h a g a r ; light brown t o strong brown on glycerol-asparagine a g a r and yeast-malt agar). Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r and tryptone-yeast broth. No pigment other than t r a c e of yellow is found in the medium i n yeast-malt agar, oatm e a l a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: 2-Glucose, _L-arabinose, sucrose, _D-xyloae,
358
i-inositol,
growth.
_D-mannitol, _D-fructose and rhamnose a r e utilized for R e p o r t s v a r y on utilization of raffinose.
Streptomyce s phaeopurpureus Shinobu. Description: Shinobu 1957, 63-67. Type strain: 146 ISP 5125 f r o m R. Shinobu a s 146. ISP description by Group B - 9 . Spore chain morphology: Section Rectiflexibiles (Fig. 150). A e r i a l mycelium is usually poorly developed o r a b s e n t on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Mature s p o r e chains m a y be observed on other media including Gauze's medium No. 1 where chains of LO to 50 o r m o r e s p o r e s may be formed. Spore surface: Smooth (Fig. 151). Color of colony: A e r i a l m a s s color probably i n the Gray o r Red c o l o r - s e r i e s , but a e r i a l sporulation on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r i s usually inadequate f o r a c c u r a t e s p o r e m a s s color determination. R e v e r s e side of colony: Yellow to yellow brown is modified by r e d (light brown, strong brown, o r g r a y i s h r e d d i s h brown on oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r ; moderate to strong reddish brown on y e a s t - m a l t a g a r ) . Substrate mycelium pigment i s not a pH indicator (or i s changed only slightly f r o m brown to yellow brown by 0.05 N HCl). Color in medium: Melanoid pigments a r e f o r m e d in peptone-yeasti r o n a g a r , tyrosine a g a r a n d tryptone-yeast broth. Yellow, orange, o r r e d pigment is found i n the medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . This pigment i s slightly pH sensitive changing f r o m reddish orange o r brown to yellow brown with 0.05 N HC1. Ca rbon u t i Lization: _D -G Luco se, _L-arabinose, s u c r o s e , _D-xy1ose, i -inositol, _D-mannitol, _D-fructose, rhamnose and raffinose a r e utilized f o r growth.
e.).
Streptomyces phaeoviridis Shinobu.
Description: Shinobu 1957, 67-
7 3 . Type strain: 503 k d . ) . ISP 5285 f r o m A. Seino, Kakken Kagaku G o . , Tokyo a s 503. ISP description by Group B -12.
Spore chain morphology: Section Rectiflexibiles (Fig. 152). Mature s p o r e chains generally long with LO to 50 s p o r e s p e r chain. This morphology is s e e n on y e a s t - m a l t a g a r , oatmeal a g a r a n d s a l t s - s t a r c h a g a r , but a e r i a l mycelium with rnature s p o r e chains i s not f o r m e d on glyceroL-asparagine a g a r . Spore surface: Smooth (Fig. 153). Color of CoLony: A e r i a l m a s s color i n the G r a y color-series on yeast-malt a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r . (White a l s o may be seen on these media a n d on the poorly sporulated s u r f a c e of glycerolasparagine agar. ) R e v e r s e side of colony: Yellow to yellow brown is modified slightly by r e d o r orange (light yellowish brown to strong brown, yellowish pink, o r g r a y i s h reddish orange). Substrate mycelium pigment is not a pH indicator. Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r a n d tryptone-yeast broth. Orange o r reddish pigment i s found in the medium i n yeast-malt a g a r , oatmeal a g a r and glycerol-asparagine a g a r . This pigment i s not pH sensitive when tested with 0.05 N NaOH o r HC1.
Figure 148. S. phaeochromogenes. spore chains. (a) (X 40p) on yeast-ma& agar, 17 days. 31 (b) (X BOO) on glycerol-asparagine agar, 14 days? Figure 149. S. phaeochromogenes. Smooth spores; electron micrograph from 2 1 Jay culture on yeast-malt agar.l Figure 150. 2. phaeopurpureus. L o n g s spore chains (X 1100) on yeast-malt agar, 21 days. Figure 1 5 1 , S. phaeopurpureus. Smooth spores; electron micrograph from 21 Zay culture on yeast-malt agar.5
360
i -inositol,
Carbon utilization: _D-Glucose, I=-arabinose, sucrose, _D-xylose, _D-mannitol, 2 - f r u c t o s e , rhamnose and raffinose a r e utilized f o r growth. S t r e ptomyces platensis Pittenger and Cottlieb. Descriptions: McGuire, B r i t i s h P a t . Specification 713, 795, Jan. 31, 19%; P i t t e n g e r and Cottlieb in Waksman 1961, 258-259. Type strain: M5-5353 (Pittenger, p e r s o n a l communication). ISP 5041 f r o m R. C. Pittenger a s M5-5353. ISP description by Group B-4. Spore chain morphology: Section S p i r a l e s (Fig. 154). Mature s p o r e chains generally LO to 50, o r s o m e t i m e s m o r e than 50, s p o r e s p e r chain. This morphology i s seen on yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 155). Color of colony: A e r i a l m a s s color i n the G r a y c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r agine a g a r . Special morphological c h a r a c t e r i s t i c : S p o r e s frequently coalesce to f o r m black, m o i s t (hygroscopic) m a s s e s of s p o r e s (Fig. 156). Sometimes much of the a e r i a l growth will be converted to a black, moist surface. R e v e r s e side of colony: G r a y i s h yellow on oatmeal a g a r and s a l t s s t a r c h a g a r ; s o m e t i m e s modified by r e d to g r a y i s h yellowish pink o r light reddish brown on y e a s t - m a l t a g a r a n d glycerol-asparagine a g a r . Reddish s u b s t r a t e mycelium pigment m a y be changed to c o l o r l e s s o r pale yellow by 0.05 N HC1. Color in medium: Melanoid pigments a r e not f o r m e d i n peptoney e a s t - i r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. T r a c e s of r e d o r yellow pigments a r e found i n the medium i n y e a s t - m a l t a g a r , oatm e a t a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . The r e d dish pigment i s somewhat pH sensitive, changing f r o m grayed r e d to nearly c o l o r l e s s with 0.05 N HC1. Carbon utilization: 2 - G lucose, s u c r o s e , i - i n o s i t o 1, _D -mannito 1, D-fructose a n d raffinose a r e utilized f o r growth, Utilization of L-arabinose and _D-xylose i s doubtful. No growth o r only t r a c e of growth with rhamnose. S t r eptomyce s pseudogri s e o lu s Okami and Ume zawa Description : Okami and Umezawa& Okami, Utahara, Oyagi, Nakamura and Umezawa. 1955, 127-128. Type s t r a i n : 534 ( x d . ) . ISP 5026 f r o m Y. Okami a s H-16C = 534 = NIHJ 224. ISP description by Group B-2. Spore chain morphology: Section S p i r a l e s (Fig. 157). Spore chains suggesting morphology m a y a l s o be common. Mature s p o r e chains moderately short, often with 3 to 10 s p o r e s p e r chain, but longer chains a r e f o r m e d on suitable media. This morphology i s seen on yeast-mzlt a g a r and s a l t s - s t a r c h a g a r but a e r i a l mycelium i s poorly developed on oatmeal a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 158). Color of colony: A e r i a l m a s s color i n the White c o l o r - s e r i e s on y e a s t - m a t t a g a r a n d s a l t s - s t a r c h a g a r ; poor sporulation on oatmeal a g a r and glycerol-asparagine a g a r . The a e r i a l mycelium is d e s c r i b e d a s g r a y i s h buff i n the original description by Okami and Umezawa (1955).
F i g u r e 152. 2. phaeoviridis. R F spore chains: (a) (X 485) on oatmeal a g a r , 14 days. 37 (b) ( X 1200) on s a l t s - s t a r c h a g a r . F i g u r e 153. 2. phaeoviridis. Smooth spores; electron micrograph f r o m 14 day culture on oatmeal a g a r . F i g u r e 154. 2. platensis. S p i r a l s p o r e chains (X 800) on glycerolasparagine a g a r , 14 days. 27 F i g u r e 155. S. platensis. Smooth spores; electron micrograph f r o m 7 day cztur-s-starch a g a r . 31 Tangled m a s s e s of s p i r a l s a r e associated with hygroscopic droplets. platensis. Hygroscopic droplets containing coalesced F i g u r e 156. m a s e e s of spores. (a) (X 400) on yeast-malt agar, 18 days (cf.Fig. 155). (b) (X 400)2m oatmeal agar, 18 days. 31 (c) (X 200) on oatmeal a g a r , 14 days.
2.
362
R e v e r s e side of colony: No distinctive pigments ( c o l o r l e s s to g r a y i s h yellow o r yellowish brown) on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r a n d glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e not f o r m e d i n peptone-yeast. i r o n a g a r , t y r o s i n e a g a r o r tryptone-yeast broth. N o pigment, o r only t r a c e of yellow, is found i n the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-GLucose, &-arabinose, _D-xylose, I-inositot, D-mannitol, ._D-fructose and rhamnose a r e utilized f o r growth. No growth o r only trace of growth with s u c r o s e and r a f f b o s e .
Actinomyces pseudolavendulae Kuchaeva, Krasil'nikov, Taptikova and Gesheva. Description: Kuchaeva e t a t . 1962, 103-124. Type s t r a i n : USSR RIA-744 = IMRU 3440 ; Krasil'nikov, p e r s o n a l cornmunication). ISP 5213 f r o m N . A . Krasillnikov a s EUA 744 = IMRU 3440. ISP description by Group B-LO. R e m a r k s : The c h a r a c t e r i z a t i o n by ISP study-group B-LO f o r Actinom y c e s pseudolavendulae Kuchaeva e t at. , ISP s t r a i n 5213 is identical i n a l l d e t a i l s with the c h a r a c t e r i z a t i o n by study-group A-6 f o r Streptom y c e s lavendulae (Waksman and C u r t i s ) Waksrnan and Henrici, ISP 5069. The c h a r a c t e r i z a t i o n f o r Streptomyces lavendulae, LSP 5069 was r e p o r t e d i n o u r previous p a p e r (Shirling and Gottlieb 1968, 138). A personal communication f r o m N.A. Krasil'nikov (April, 1964) gives IMRU 3440 = RLA-744 a s the type s t r a i n f o r Actinomyces pseudolavendulae. IMRU 3440 is given a s the type s t r a i n f o r Streptomyces Lavendulae by Waksman (1961) and was used a s the type s t r a i n f o r the ISP study of t h i s species. Actinomyces pseudolavendutae Kuchaeva et at. 1962 i s r e g a r d e d h e r e a s a n objective synonym f o r Streptomyces Lavendulae (Waksman and C u r t i s ) Waksman and Henrici 1948; and ISP s t r a i n 5213 is equivalent to ISP 5069 Streptomyces Lavendulae 9 . ~
w.
Actinomyces pseudovenexue lae Kuchaeva, K r a sillnikov, Taptikova 1962, 103-124. Type strain: and Gesheva. Description: Kuchaeva USSR RIA-742 = IMRU 3774 (ibid. ; Kraeitlnikov, p e r s o n a l communication). ISP 5212 f r o m N.A. Krasil'nikov a s RIA-742 = IMRU 3774. ISP description by Group B-9. Spore chain morphology: Section Rectiflexibiles. Mature s p o r e chains generally long, often with m o r e than 50 s p o r e s p e r chain (Fig. 159). This morphology is s e e n on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r a n d glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 160). Color of colony: A e r i a l m a s s color i n the Red c o l o r - s e r i e s (5dc, g r a y i s h yellowish pink) on y e a s t - m a l t a g a r , o a t m e a l a g a r a n d s a l t s s t a r c h a g a r ; G r a y o r White c o l o r - s e r i e s on glycerol-asparagine a g a r . (Gray o r W h i t e is a l s o s o m e t i m e s r e p o r t e d f o r y e a s t - m a l t a g a r , oatmeal agar and ealte-starch agar.) R e v e r s e side of colony: No distinctive pigments (light g r a y i s h yellow to orange yellow o r Light yellowish brown) on y e a s t - m a l t a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n aga-r, tyrosine a g a r and tryptone-yeast broth. T r a c e of grayish yellow pigment may be found in the medium in yeast-malt a g a r , oatmeal a g a r , s a l t a - a t a r c h a g a r and glycerol-asparagine a g a r . This pigment is
s.
Figure 157. S. pseudogriseolus. Spiral spore chains (X 380) on Waksman's starch agar B, 21 days. Figure 158. 5. seudogriseolus. Smooth spores; electron micrograph from 21 day'culture on yeast-malt agar. p 9 pseudovenezuelae. spore chains (X 1100) on Figure 159. glycerol-asparagine agar, 14 days. Figure 160. A. pseudovenezuelae. Smooth spores; electron micrograph from 14 xay culture on glycerol-asparagine agar.
'*
A.
RF
'
364
not pH sensitive when tested with 0.05 N NaOH o r HC1. Carbon uti lization: _D-Gluco se, &-arabinose, sucrose, _D -xy lo s e , i -inositol, _D-mannitol, _D-fructose, rhamnose and raffinose a r e utilized for growth.
Streptomycee purpeofuscus Yamaguchi and Saburi. Description: Yamaguchi and Saburi 1955, 203-207. Type strain: H-5080 &d2). ISP 5283 f r o m T. Yamaguchi a s H-5080. ISP description by Group B -12. Spore chain morphology: Section Rectif lexibiles (Fig. 161). Mature spore chains generally long, often with m o r e than 50 s p o r e s p e r chain. This morphology i s seen on yeast-malt agar, oatmeal agar, s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 162). Color of colony: Aerial m a s s color in the Gray c o l o r - s e r i e s on yeastmalt agar, oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Reverse side of colony: Distinctive pigments a r e usually not produced (grayish yellow to olive brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r o r glycerol-asparagine a g a r , but d a r k brown to reddish brown substrate mycelium pigment may be produced. The original description (Yamaguchi and Saburi, 1955) d e s c r i b e s purple to brown r e v e r s e mycelium color but notes that this coloration may be lost on repeated t r a n s f e r . Color in medium: Melanoid pigments a r e formed in peptone-yeasti r o n a g a r , but only a s t r a c e s of brownish black color in tyrosine a g a r o r tryptone-yeast broth. T r a c e s of red, violet o r yellow pigment may be found in the medium i n yeast-malt a g a r , oatmeal a g a r and s a l t s s t a r c h a g a r . This pigment is not pH sensitive when tested with 0.05 N NaOH o r HCl. Carbon utilization: _D-Glucose, 2 - a r a b i n o s e and _D-xylose a r e utilized f o r growth. Utilization of _D-fructose i s doubtful. No growth o r only t r a c e of growth with sucrose, i-inositol, _D-maanitcl, rhamnose and raffinose. Streptomyces pyridomyceticus Okami and Umezawa. Descriptions and nomenclature: Tentative name with incomplete and tentative d e s criptinn in Maeda, Kosaka, Okami and Umezawa 1953, 140; Streptomyces albidofuscus Okami and Umezawa i n Okami, Maeda and Umezawa 1954, 55-56; authors' rejection of Streptomyces albidofuscus with assignment of the name Streptomyces pyridomyceticus Okami and Umezawa i n Umezawa and Okami 1957, 172. Type strain: 451-A8 (Okami and 5 ) . ISP 5024 f r o m Y . U m e z a w a h Okami, Maeda and Umezawa 1954, 5 Okami as 451-A8 = NIHJ 284. ISP description by Group B-3. Spore chain morphology: Section not determined (possibly Spirales o r RF). Mature spore chains a r e generally difficult to find on yeastmalt agar, oatmeal a g a r , s a l t s - s t a r c h a g a r , glycerol-asparagine a g a r and other ISP media a s well as on Hickey and T r e s n e r ' s amidex a g a r . R a r e spore chains were short with 3 to LO s p o r e s p e r chain and were generally twisted or curved into hooks resembling incomplete s p i r a l s . (Fig. 163), although spore chains were a l s o sometimes found (Fig. 164). The original descriptions (Maeda e t at., 1963 and Okami and Umezawa, 1954) a l s o note the general absence of good a e r i a l mycelium on most media. In the original description loose s p i r a l s a r e described a s occurring on the s p a r s e a e r i a l mycelium of synthetic media. Spore surface: Smooth (Fig. 165). Spores were i r r e g u l a r in size and w e r e often i n t e r s p e r s e d with imperfect spores. This c h a r a c t e r i s t i c i s a l s o
F i g u r e 161. 2. purpeofuscus. spore chains on yeast-malt agar, 21 days (a, X 300; b, X 1200).23 F i g u r e 162. 2. purpeofuscus. Smooth spores; electron micrograph f r o m 14 day culture on oatmeal a g a r . 22 F i g u r e 163. 2. pyridomyceticus. Isolated spore chains forming hooks, incomplete s p i r a l s o r s p i r a l s of only one o r two t u r n s (X 600): (a) On glycerol-asparagine agar, 14 days; (b, c) On yeast-malt agar, 14 days; (d) On glycerol-asparagine agar, 18 days. 2o Aerial mycelium i s difficult t o find on this strain. spore chains (X 600) on glycerolF i g u r e 164. S. pyridomyceticus. asparagine agar, 12 days ( s e e also, F i g u r e 163).20 F i g u r e 165. S. pyridomyceticus. Smooth spores; electron micrograph f r o m 14 z a y culture on yeast-malt a g a r . lo
366
INTERNATIONAL
JOURNAL
mentioned i n the f i r s t description. Color of colony: A e r i a l mycelium is poorly developed o r absent on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r a gine a g a r a s well a s on other ISP media. When p r e s e n t on y e a s t - m a l t a g a r o r oatmeal a g a r , a e r i a l mycelium i s thin a n d white. The original description d e s c r i b e s the a e r i a l mycelium a s thin a n d white, s o m e t i m e s with g r a y o r brownish gray tinge on Czapek's a g a r . R e v e r s e side of colony: No distinctive pigments (colorless o r g r a y i s h yellow o r olive brown) on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r . Color in medium: Melanoid pigments a r e not f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. N o pigment i s found i n the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: No growth o r only t r a c e of growth with 2-Gtucose, L -a ra bin0 s e , s u c r o s e , _D -xy lo s e , L-ino s i t o 1, _D -manni t o 1, , D f i'xto s e , rhamnose o r raffinose. Absence of growth with _D-gtucose and a l l other carbon s o u r c e s tested, suggests that P r i d h a m and Gottlieb's b a s a l medium will not support growth of this strain. The original description (Okami and Umezawa 1954) indicates that _D-glucose, &-arabinose, s u c r o s e , 2 - x y l o s e and _D-fructose can be utilized f o r growth with Caapek's a g a r a s the b a s a l medium.
S t r eptomyce s racemochromogenus Sugai. Description: Sugai 1956, 170-179. Type strain: 229 a d . ) . ISP 5194 f r o m M. Oda, Meiji Seika Go. ISP description by Group B-4. Spore chain morphology: Section Retinaculiaperti including many R F s p o r e chains (Fig. 166a), and s o m e long chains with t e r m i n a l hooks o r primitive s p i r a l s of wide d i a m e t e r (Fig. 166b). Mature s p o r e chains generally long with LO t o 50, o r often m o r e than 50, s p o r e s p e r chain. This morphology is s e e n on y e a s t - m a t t a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 167). Color of colony: A e r i a l m a s s color i n the Red c o l o r - s e r i e s (5ec a n d 5cb, g r a y i s h yellowish pink) on yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r . A r e a s of g r a y o r white sporulation m a y a l s o be seen. R e v e r s e side of colony: N o distinctive pigments on oatmeal a g a r , but y e l l w t o yellow browp s u b s t r a t e mycelium color may be modified i n some a r e a s by blue pigment (dark olive green, g r a y i s h g r e e n o r grayish blue) on y e a s t - m a l t a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . The blue pigment is not always present. When p r e s e n t , i t can be intensified f r o m g r a y i s h g r e e n to grayish blue o r violet by 0.05 N NaOH o r f r o m g r a y g r e e n to reddish g r a y by 0.05 N HC1. Color in medium: Melanoid pigments a r e produced i n peptone-yeasti r o n a g a r , tyrosine a g a r and tryptone-yeast broth. No pigment other than a t r a c e of yellow i s found in the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Carbon utilization: _D-Glucose, 5 - a r a b i n o s e and s u c r o s e a r e utilized -inositol, f o r growth. Only a t r a c e of growth is found on _D-xylose, Q-mannitot, _D-fructose, rhamnose and raffinose.
morphology including both F i g u r e 166. 2. racemochromogenus. straight spore chains and terminal spirals: (a) R.F chains (X 400) on yeast-malt a g a r , 17 days. 31 (b) T y p i c a l 2 terminal hooks and s p i r a l s (X 200) on oatmeal agar, 14 days." F i g u r e 167. 2. racemochromogenus. Smooth spores; electron micrograph f r o m 14 day culture on oatmeal agar. 40 F i g u r e 168. S. rimosus. Spiral spore chains (X 1100) from,14 day cultures;n:past-malt agar, (b) oatmeal agar, (c) glycerolasparagine agar. F i g u r e 169. 3.rimosus. Smooth spores; electron micrograph f r o m 14 day culture on yeast-malt agar.'
368
INTERNATIONAL
JOURNAL
Streptomyces r i m o s u s (Sobin, Finlay and Kane) Waksman. Descriptions: Sobin, Finlay a n d Kane, U.S. P a t . 2,516,080 July 18, 1950. Waksman & W a k s k n a n d Lechevalier 1953, 47. Type -strain: P f i z e r F.D. 10326 = NRRL 2234, IMRU 3558, ATCC 10970 (Routien, personal communication). ISP 5260 f r o m J.B. Routien, Chas. P f i z e r and Co. , Inc. a s F D 10326. ISP description by Group B-9. Spore chain morphology: Section S p i r a l e s (Fig. 168), but many i m p e r f e c t s p i r a l s , hooks, and Loops suggesting morphology a r e a l s o p r e s e n t . Mature s p o r e chains s h o r t to moderately long with 3 to LO, o r often m o r e than LO, s p o r e s p e r chain. This morphology i s seen on y e a s t - m a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r , glycerol-asparagine a g a r and on carbon utilization medium plus r)-glucose. Spore surface: Smooth (Fig. 169). Color of colony: A e r i a l m a s s color i n the Red o r White c o l o r - s e r i e s on yeast-malt a g a r and in the Yellow o r White c o l o r - s e r i e s on o a t m e a l a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Representative color tabs a r e 3ca (pale orange yellow), 5cb (grayish yellowish pink) and 2ca (pate yetlow) f r o m the Red color-wheel; 2db and 2ba (pale yellow) f r o m the Yellow color-wheel; and a (white) f r o m the White c o l o r wheel. R e v e r s e s i d e of colony: G r a y i s h yellow on oatmeal a g a r and s a l t s s t a r c h a g a r ; g r a y i s h yellow modified slightly by r e d ( m o d e r a t e to d a r k orange-yellow) on y e a s t - m a l t a g a r and glycerol-asparagine a g a r . Subs t r a t e mycelium pigment is not a pH indicator. Color in medium: Melanoid pigments a r e not f o r m e d i n peptoneyeast-iron a g a r , tyrosine a g a r o r tryptone-yeast broth. T r a c e of yellow pigment is found in the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . This pigment is not pH sensitive when tested with 0.05 N NaOH o r H C l . mannitol, Carbon utilization: _D-Gtucose, &-arabinose, i-inositol, 2D-fructose and raffiaoee a r e utilized f o r growth. Utilization of _Dxylose i s doubtful. No growth o r only t r a c e of growth with s u c r o s e and raffinose.
Streptomyces rochei B e r g e r , Jampolsky and Goldberg. Description: Name only, without description i s found i n B e r g e r , Jampolsky and Coldberg 1949, 476-478. Description i s given i n Waksman and Lechevalier a_l. 1957, 133 1953, 40-41 and by Preobrazhenskaya Et 51. & Gauze (Engl. t r a n s l . , 106). Type strain: X 15 ( B e r g e r , personal communication). ISP 5231 f r o m J. B e r g e r , Hoffman-LaRoche, Inc. as X 15. ISP description by Group B-8. Spore chain morphology: Section S p i r a l e s (Fig. 170a). S p i r a l s a r e usually open, s o m e t i m e s a l m o s t flexuous (Fig. 170 b, c). Mature s p o r e chains moderately tong with LO to 50, o r often m o r e than 50 s p o r e s p e r chain. This morphology is s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth to warty (Fig. 171). Color of colony: A e r i a l m a s s color in the G r a y c o l o r - s e r i e s on yeastm a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine agar. R e v e r s e side of colony: No distinctive pigments (grayish yellow to yellowish brown) on y e a s t - m a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r .
c t
Figure 170. 2. rochei. (a) Spiral spore chains (X 300) on glycerolasparagine agar, 7 d a y s T b , c) Open spirals to flexuous chains (X 400) on oatmeal agar, 14 days. 41 Figure 171. 5. rochei. Smooth to warty spores; electron micrograph from 14 day culture on oatmeal agar. l 9
370
INTERNATIONAL
JOURNAL
Color in medium: Melanoid pigments a r e not f o r m e d in peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. N o pigment i s found in the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-GLucose, _L-arabino se, 2 - x y l o s e , i -inositol, D-mannitol, _D-fructose and rhamnose a r e utilized f o r growth. No growth or only t r a c e of growth on s u c r o s e and raffinose. Streptomyces roseo-luteus B e s s e t t . Description: B e s s e l l , U.S. Patent 3,013, 947 Dec. 19, 1961). Type s t r a i n : NRRL 2776 = NCIB ISP 5240 f r o m C. W. Hesseltine (NRRL) a s NRRL 2776. 8984 (ifi.). ISP description by Group B-6. Spore chain morphology: Section S g i r a t e s . Tight s p i r a l s of one to many t u r n s occur at ends of long s p o r e chains (Fig. 172). These s o m e t i m e s s u g g e s t R A morphology. Mature s p o r e chains generally long with LO to 50, o r often m o r e than 50, s p o r e s p e r chain. This morphology i s seen on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerola s p a r a g i n e a g a r . Spore surface: Smooth (Fig. 173). Color of colony: A e r i a l m a s s color i n the Red c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r agine a g a r . One o b s e r v e r placed this s t r a i n i n the G r a y c o l o r - s e r i e s with color tab 5fe (light g r a y i s h reddish brown) a s the n e a r e s t matching color f r o m the G r a y color wheel. N e a r e s t matching t a b s i n the Red c o l o r - s e r i e s w e r e 5ge and 4ge (light grayish reddish brown). R e v e r s e side of colony: Yellow to orange yellow on y e a s t - m a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ; s u b s t r a t e mycelium pigment i s not a pH indicator. Color in medium: Melanoid pigments a r e not f o r m e d in peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. Yellow pigment i s found in the medium in yeast-matt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . This pigment is not pH sensitive when tested with 0.05 N NaOH o r HC1. Carbon utilization: _D-GLucose, _D-xylose, _D-mannitol, _D-fructose and raffinose a r e utilized f o r growth. No growth o r only t r a c e of growth with _L-arabinose, s u c r o s e , 1-inositol and rhamnose. Streptomyces r o s e o s p o r u s F a l c z o d e Morais and Dalid Maia. Description: Streptomyces venezuelae v a r . r o s e o s p o r i F a l c z o d e Morais, Dalid h4aia and Genn 1958, 102-103. Change in name (rank): Streptomyces r o s e o s p o r u s Morais and D. Maia ( , , c , nom nov 1961, 43. Type strain: 4192 (Falcxo d e Morais and Dalid Maia 1961, 43). ISP 5122 f r o m J.O. FalcZo d e Morais a s 4192. ISP description by Group B - 3 . Spore chain morphology: Section Rectiflexibiles. Mature s p o r e chains generally long, often with m o r e than 50 s p o r e s p e r chain (Fig. 174). This morphology i s seen on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 175). Color of colony: Aerial m a s s color i n the Red c o l o r - s e r i e s on y e a s t m a l t a g a r , oatmeal a g a r and s a l t s - s t a r c h a g a r . (White, Yellow o r Red c o l o r - s e r i e s on glycerol-asparagine a g a r . ) R e v e r s e side of colony: No distinctive pigments (grayish yellow to yellowish brown) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r .
F i g u r e 172. S. roseo-luteus. Spiral spore chains ( X 500) on s a l t s - s t a r c h agar, 3 X a y s 7 F i g u r e 173. 2. roseo-luteus. Smooth s p o r e s (Au/Pd shadowed); electron micrograph f r o m 21 day culture on s a l t s - s t a r c h agar. 44 F spore chains. (a) ( X 300) on oatmeal F i g u r e 174. S . roseosporus. R agar, 14 days. lo (b) (X 600) on glycerol-asparagine a g a r , 21 days. 2o F i g u r e 175. 2. roseosporus. Smooth spores; electron micrograph f r o m 14 day culture on s a l t s - s t a r c h a g a r .
372
INTERNATIONAL
JOURNAL
Color in medium: Melanoid pigments a r e not f o r m e d i n peptoneyeast-iron a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment i s found in the medium in yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _DGlucose, ,L -arabinose, 2 - x y l o s e and r h a m n o s e a r e utilized f o r growth. Utilization of _D-fructose i s doubtful. No growth o r only t r a c e of growth with s u c r o s e , t -inositol, D-mannitol aad raffinose. Actinomyces roseoviridis Preobrazhenskaya. Description: P r e o brazhenskaya @ Gauze 51. 1957, 57. Type strain: 3617 ( P r e o b r a z h e n skaya, personal communication; G. F. Gauze, cited by Gottlieb 1968, 20). Streptomyces roseoviridis (Preobrazhenskaya) P r i d h a m e t a l . 1958, 61. ISP 5175 f r o m T . P . Preobrazhenskaya a s 3617. ISP description by Group A-12. Spore chain morphology: Section Rectiflexibiles (Fig. 176). Mature s p o r e chains generally LO to 50 s p o r e s p e r chain. This morphology is s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerola s p a r a g i n e a g a r . Spore surface: Smooth (Fig. 177). Color of colony: A e r i a l m a s s color i n the Red o r Yellow c o l o r - s e r i e s on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerola sparagine a g a r . R e v e r s e side of colony: No distinctive pigment on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r and tryptone-yeast broth. P i g m e n t s other than melanoids a r e not f o r m e d in y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-GLucose, _L-arabinose a n d _D-xylose a r e utilized f o r growth. No growth o r only t r a c e of growth on s u c r o s e , -inositol, D-mannitol, rhamnose and raffinose. Utilization of _D-fructose i s doubtful.
c t
Actinomyces rubiginosohelvolus Kudrina. Description: Kudrina in Gauze et a l . 1957, 78 and 89. Type s t r a i n : L0/53 k d . ; s e e a l s o Gauze i & Cottlieb 1968). Streptomyces rubiginosohelvolus (Kudrina) P r i d h a m . e t at. 1958, 59. ISP 5176 f r o m T . P . Preobrazhenskaya a s 10/53. ISP description by Group A-12. Spore chain morphology: Section Rectiflexibiles (Fig. 178). A e r i a l mycelium i s s o m e t i m e s poorly developed on ISP media. Mature s p o r e chains, when formed, contain LO to 50 o r m o r e s p o r e s p e r chain on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r agine a g a r . Spore surface: Smooth (Fig. 179). Color of colony: A e r i a l m a s s color in the Yellow c o l o r - s e r i e s on y e a s t - m a l t a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . A e r i a l mycelium is poorly developed on oatmeal a g a r . One o b s e r v e r a l s o r e c o r d e d tabs 2ca(pale yellow) and 3ca (pale orange yellow) f r o m the Red c o l o r - s e r i e s a s a second color on y e a s t - m a l t a g a r a n d glycerolasparagine agar. R e v e r s e side of colony: Colorless to pale yellow o r grayed yellow on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerola s p a r a g i n e a g a r . However, yellowish r e v e r s e pigment is a pH indicat o r changing f r o m yellow to pink with addition 0.05 N NaOH.
Figure 176. A. roseoviridis. RF spore chains: (a) (X 240) on oatmeal agar, 14-days. (b) ( X 400)on yeast-malt agar, 21 days. Figure 177. A. roseoviridis. Smooth spores; electron micrograph from 21 day c & u r e on yeast-malt agar. 23 F spore chains (X 400) on yeastFigure 178. _A. rubiginosohelvolus. R malt agar, 21 days.z3 Figure 179. _A. rubiginosohelvolus. Smooth spores; electron micrograph from 21 day culture on yeast-malt agar. 23
374
Color i n medium: Melanoid pigments not f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. N o pigments o r only t r a c e of yellow pigment found in yeast-malt a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r o r glycerol-asparagine a g a r . o s e , _D mannito 1, Carbon uti t i zation : _D -G luc o s e, 4 -a rabino se , ,D -xy L D-fructose and rhamnose a r e utilized f o r growth. No growth o r only t r a c e of growth on sucrose, i - i n o s i t o l and raffinose.
Actinomyces rubiginosus Preobrazhenskaya, B linov and Ryabova. Description: Preobrazhenskaya et al.in Gauze e t a t . 1957, 132 and 134. Type s t r a i n : 11852 (Gauze & Cotttieb, 1968). Streptomyces rubiginosus (Preobrazhenskaya, Blinov and Ryabova) P r i d h a m e t a l . 1958, 70. I S P 5177 f r o m T.P. Preobrazhenskaya a s 11852. ISP description by Group A-12. Spore chain morphology: Section Spirales. S p i r a l s a r e b e s t developed on s a l t s - s t a r c h a g a r . A e r i a l mycelium and s p o r e chains m a y be poorly developed and atypical on y e a s t - m a l t a g a r , oatmeal a g a r and glycerota s p a r a g i n e a g a r (Fig. 180). Mature s p o r e chains generally LO to 50 s p o r e s p e r chain. Spore surface: Spiny (Fig. 181). f colony: A e r i a l mass color i n the G r a y c o l o r - s e r i e s on Color o y e a s t - m a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r agine a g a r . R e v e r s e side of colony: No distinctive pigment (yellowish brown, grayed orange brown o r yellowish gray) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Color in medium: Melanoid pigments not f o r m e d i n peptone y e a s t i r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment is found i n medium i n yeast-malt a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . T h e r e may be a t r a c e of r e d soluble pigment i n oatmeal a g a r ; this pigment i s not pH sensitive. Carbon utilization: _D-Glucose, s u c r o s e , _D-xylose, i -inositol, D-mannitol, _D-fructose and rhamnose a r e utilized f o r growth. Growth on s u c r o s e i s l e s s abundant than on other carbon s o u r c e s . N o growth o r only t r a c e of growth on raffinose.
Streptomyces s c a b i e s (Thaxter) Waksman and Henrici. Descriptions: Oospora s c a b i e s Thaxter 1892, 159-160. Actinomyces s c a b i e s (Thaxter) Giissow 1914, 433; Millard and B u r r 1926, 628- 629. Streptomyces s c a b i e s (Thaxter) Waksman a n d Henrici 1948, 957. Type strain: IMRU 3018 (Waksman 1961, 274-275). ISP 5078 f r o m S.A. Waksman a s IMRU 3018. ISP description by Group B -4. Spore chain morphology: Section Rectiflexibiles (Fig. 182). Mature s p o r e chains moderately s h o r t with 3 to LO, o r s o m e t i m e s m o r e than 10, s p o r e s p e r chain. This morphology i s seen on y e a s t - m a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 183). Color of colony: A e r i a l m a s s color in the G r a y c o l o r - s e r i e s on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine agar. R e v e r s e side of colony: Yellow to yellow brown is usually modified by g r e e n to light g r a y i s h olive, olive, o r olive brown on y e a s t - m a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Substrate mycelium pigment i s not a pH indicator.
F i g u r e 180. A. rubiginoeus. Spiral spore chains; imperfect s p i r a l s (X 400) on yeast-malt agar, 21 days." F i g u r e 181. _A. rubiginosus. Spiny spores; electron micrograph f r o m 21 day culture on oatmeal agar. F i g u r e 182. S. scabies. RF epore chains: (a) (X 400) on yeast-malt agar, 17-days. 31 X ( ) b 800) on s a l t s - s t a r c h agar, 14 days. F i g u r e 183. 5. scabies. Smooth spores; electron micrograph f r o m 14 day culture on oatmeal a g a r . F i g u r e 184. S. sindenensis. R F spore chains (X 200) on oatmeal a g a r , 14 days.m F i g u r e 185. sindenensis. Long chains of smooth spores; electron micrograph (X 2700) f r o m 16 day culture on yeast-malt agar. F i g u r e 186. 2. sindenensis. Smooth s p o r e s (X 20,000); electron micrograph f r o m 14 day culture on yeast-malt agar.
''
''
''
2.
''
376
INTERNATIONAL
JOURNAL
Color i n medium: Melanoid pigments a r e not f o r m e d i n peptoneyeast-iron a g a r , tyrosine a g a r o r tryptone-yeast broth. Yellow o r greenish yellow pigment may be found i n the medium i n y e a s t - m a l t a g a r and oatmeal a g a r , but not i n s a l t s - s t a r c h a g a r o r g l y c e r o l - a s p a r agine a g a r ; t h i s pigment is not pH sensitive when tested with 0.05 N NaOH o r H C l . Carbon utilization: 2-Glucose, &-arabinose, _D-xylo se, _D -mannitol , D - f r u c t o s e and rhamnose a r e utilized f o r growth. No growth o r only i -inositol and raffinose. t r a c e of growth with s u c r o s e , Streptomyces sindenensis Nakazawa and Fujii. Descriptions: Nakazawa and Fujii 1957, 109-110; Nakazawa 1964, 69-70. Name only, without description, i n Nakazawa, Fujii, Inoue, Hitomi, Miyake and Kaneko 1954, 168. Type s t r a i n : 1071 ( i . . ) . ISP 5255 f r o m K . Nakazawa a s 1071 = IF0 3399. ISP description by Group B -11. Spore chain morphology: Section Rectiflexibiles (Fig. 184). Mature s p o r e chains short, often with only 3 to LO s p o r e s p e r chain (longer chains may be found on i m p r e s s i o n mounts o r electron m i c r o g r a p h s . Fig. 185). This morphology i s s e e n on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 186). Color of colony: A e r i a l m a s s color i s difficult to determine because sporutating a e r i a l mycelium i s poorly developed on ISP media. When sporulation occurs, i t i s usually i n the White o r Red (5cb, grayish yellowish pink) c o l o r - s e r i e s on y e a s t - m a l t a g a r and i n the White o r yellow c o l o r - s e r i e s on oatmeal a g a r . When present, sporulating a e r i a l mycelium i s i n the White c o l o r - s e r i e s on s a l t s - s t a r c h a g a r a n d glycerolasparagine agar. R e v e r s e side of colony: No distinctive pigments (colorless o r pale grayish yellow) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g tycerol -asparagine a g a r . Color in medium: Melanoid pigments a r e not f o r m e d i n peptone-yeasti r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment i s found in the medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r g Lyce r o 1 -a spa ragine a g a r . Carbon utilization: _D-GLucose, _L-arabinose, _D-xylose, _D-mannitol and 2 - f r u c t o s e a r e utilized f o r growth. No growth o r only t r a c e of growth with s u c r o s e , i -inositol, rhamnose and raffinose. Streptomyces somaliensis (Brumpt) Waksman and Henrici. Description: Indiella somaliensis B r u m p t 1906, 555 and 561-562. See also: Actinomyces somaliensis E r i k s o n 1935, 17-18; Streptomyces somaliensis (Brumpt) Waksman and Henrici 1948, 965. Type s t r a i n : 3236 (Erikson 1935, 17, neotype). ISP 5267 f r o m S . T. Cowan, NCTC a s NCTC 3236. ISP description by Group B-11. Spore chain morphology: Section Rectiflexibiles; flexuous chains a r e m o s t common (Fig. 187). Mature s p o r e chains generally LO to 50, o r s o m e t i m e s m o r e than 50. s p o r e s p e r chain. This morphology i s seen on y e a s t - m a t t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerola s p a r a g i n e a g a r . Spore surface: Smooth (Fig. 188). Color of colony: A e r i a l m a s s color i n the Yellow c o l o r - s e r i e s (2db o r Zba, pale yellow) on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . (One o b s e r v e r selected color tab
Figure 187. S. somaliensis. & F spore chains on oatmeal agar, 14 days. 36 Figure 188. S. somaliensis. Smooth spores; electron micrograph from 14 day culture on yeast-malt agar. Figure 189. S. s heroides. Spiral spore chains (X ca 900) on saltsstarch agar, '14 days. Figure 190. spheroides. Smooth spores; electron micrograph from 14 day culture on salts-starch agar.
5.
*-
378
3ca, pale orange yellow, f r o m the Red color wheel a s the m o s t r e p r e sentative c o l o r . ) A e r i a l mycelium i s often poorly developed on much of the culture surface on ISP media. Good sporulation in the Yellow c o l o r - s e r i e s w a s observed on Czapekls a g a r . R e v e r s e side of colony: No distinctive pigments (grayish yellow to grayish g r e e n i s h yellow and light olive brown) on yeast-malt a g a r , s a l t s s t a r c h a g a r and glycerol-asparagine a g a r . Color in medium: Melanoid pigments a r e not formed in peptoney e a s t - i r o n a g a r , tyrosine a g a r o r tryptone-yeast broth. Yellow pigment is found i n the medium i n yeast-matt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r ; this pigment i s not pH sensitive when tested with 0.05 N NaOH or HC1. Carbon utilization: D-Glucose, _L-arabinose, sucrose, _D-xylose, D-mannitol and E - f r u c G s e a r e utilized f o r growth. No growth o r only t r a c e of growth with i-inositol, rhamnose o r raffinose.
Streptomyces spheroides Wa llick, Harris, Reagen, Ruger and Woodruff. Description: Wallick e t a t . 1956, 909-917. Type strain: M A 319 (Woodruff, personal communication 1964). ISP 5292 f r o m H. B . Woodruff, Merck, S h a r p and Dohme a s M A 319 ( =NRRL 2449). ISP description by Group B-12. Spore chain morphology: Section S p i r a l e s (Fig. 189). Mature s p o r e chains generally long, often with m o r e than 50 s p o r e s p e r chain, This morphology i s seen on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 190). Color of colony: A e r i a l m a s s color i n the Yellow c o l o r - s e r i e s on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . R e v e r s e side of colony: No distinctive pigments ( c o l o r l e s s to pale yellow o r light yellowish brown) on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Color in medium: Melanoid pigments a r e not forlned in peptoney e a s t - i r o n a g a r , tyrosine a g a r or tryptone-yeast broth. No pigment i s found i n the medium i n yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-Glucose, _L -arabinose, s u c r o s e , _D-xylose, D-mannitol, 2 - f r u c t o s e and rhamnose a r e utilized f o r growth. Only a t r a c e of growth is found on 'i-inositol and raffinose. A t r a c e of growth a l s o o c c u r s on the carbon-free b a s a l medium.
Actinomyce s streptomycini Krasil'nikov. Descriptions : Actinomyce s globisporus streptomycini Krasilanikov 1949, 100; Actinomyces streptomyciniKrasil'nikov 1955, 12-18; s e e a l s o Nikitina e t at. 1960, 85-103 (Engl. t r a n s l . 80-98). Type strain: RIA 333 = INh4I 1448 (Krasil'nikov, p e r s o n a l communication, 1964). ISP 5200 f r o m N.A. Krasilanikov a s RLA 333 = INMX 1448 ( =ATCC 15886). Note: This is a subjective synonym f o r Streptomyces g r i s e u s (Krainsky) Waksman and Henrici, ISP 5236 = I M R U 3463 1.v. ISP description by Group B -3. Spore c h i p morphology: Section Rectiflexibiles, flexuous chains a r e m o s t common (Fig. 191). Mature spo;e chains generally long, often with m o r e than 50 s p o r e s p e r chain. This morphology i s s e e n on yeast-malt a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r agine a g a r . Spore surface: Smooth (Fig. 192).
Figure 191. A. streptomycini. RF spore chains ( X 300) on yeast-malt agar, 15days. " Figure 192. streptomycini. Smooth spores; electron micrograph from 14 day culture on yeast-malt agar. I t Figure 193. 2. tanashiensis. RF spore chains (X 312) on glycerolasparagine agar, 14 days? Figure 194. S. tanashiensis. Smooth spores; electron micrograph from 10 day culture on oatmeal agar.
A.
380
INTERNATIONAL
JOURNAL
Color of colony: A e r i a l mass color i n the Yellow c o l o r - s e r i e s on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerola s p a r a g i n e a g a r ( s o m e t i m e s i n the White c o l o r - s e r i e s on glycerola sparagine a g a r ) . R e v e r s e side of colony: No distinctive pigments (colorless, grayish yellow o r light olive brown) on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r a n d glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e not f o r m e d i n peptone-yeasti r o n a g a r , t y r o s i n e a g a r o r tryptone-yeast broth. N o pigment i s found in the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r o r g lycerol-asparagine a g a r . Carbon uti Liza tion : _O -G lu c o se, _D -xy lose, _D ma nnit o 1 and _D f r u c to s e and utilized f o r growth. No growth o r only t r a c e of growth with 4arabinose, s u c r o s e , i-inositol, rhamnose and raffinose.
Streptomyces tanashiensis Hata, Ohki and Higuchi. Descriptions: Hata, O h k i a n d Higuchi 1952a, 529-534; 1%2b, 67-77 (448-457); 1952c, 188-189. Type s t r a i n : 144 (ibid.). ISP 5195 f r o m Y. Yagisawa f r o m T. Hata, Kitasato Inst., a s 144. ISP description by Group B-5. Spore chain morphology: Section Rectiflexibiles (Fig. 193). Mature s p o r e chains generally LO to.50, o r s o m e t i m e s m o r e than 50, s p o r e s p e r chain. This morphology i s s e e n on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 194). Color of colony: A e r i a l m a s s color i n the G r a y c o l o r - s e r i e s on y e a s t - m a l t a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r and g l y c e r o l - a s p a r a gine a g a r . R e v e r s e side of colony: No distinctive pigments (pale yellow to light olive brown) on y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r and tryptone-yeast broth, but not on tyrosine a g a r . No pigment is found i n the medium i n y e a s t - m a l t a g a r , oatmeal a g a r , s a l t s s t a r c h a g a r o r glycerol-asparagine a g a r . Carbon utilization: _D-Glucose, _L-arabinose and _D-xylose a r e utilized f o r growth. R e p o r t s v a r y on utilization of _D-fructose. Only a t r a c e of growth is found with s u c r o s e , i-inositol, _D-rnannitol, r h a m n o s e and raffinose. Actinomyces violascens P r e o b r a z h e n s k a y a a n d Sveshnikova. D e s cription: P r e o b r a z h e n s k a y a a n d Sveshnikova & Gauze et at. 1957, 32 and 41. Type strain: 3959/54 (Ibid.; see a l s o Gauze&Gottlieb 1968). S t r e p t o m y c e s violascens (Preobrazhenskaya a n d Sveshnikova) S P 5183 f r o m T . P . Preobrazhenskaya a s P r i d h a m %tat. 1958, 68. I S P description by Group A-13. 3959/54. I Spore morphology: Section S p i r a l e s (Fig. 195). Mature s p o r e chains LO t o 50 o r m o r e s p o r e s p e r chain. This morphology i s s e e n on y e a s t m a l t a g a r , oatmeal a g a r , s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r . S p o r e sqface: Spiny (Fig. 196). Color of colony: A e r i a l m a s s color i n the Violet c o l o r - s e r i e s on s a l t s - s t a r c h a g a r a n d glycerol-asparagine a g a r ; Violet o r White colorseries on oatmeal a g a r ; Violet, White o r Red c o l o r - s e r i e s on y e a s t malt a g a r .
Figure 195. A. violascens. Spiral spore chains (X 500) on yeast-malt agar, 24days. l5 Figure 196. 4. violascens. Spiny spores; electron micrograph from 24 day culture on yeast-malt agar. 24
38 2
INTERNATIONAL
JOURNAL
R e v e r s e side of colony: No distinctive pigment (grayed yellow to yellow-brown) on y e a s t - m a l t a g a r , o a t m e a l a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Color i n medium: Melanoid pigments a r e f o r m e d i n peptone-yeasti r o n a g a r , t y r o s i n e a g a r and t r y p t o n e - y e a s t broth. P i g m e n t s o t h e r t h a c m e l a n o i d s not f o r m e d ( o r only t r a c e s of yellow) i n y e a s t - m a l t a g a r , oatmeal a g a r , salts-starch a g a r and glycerol-asparagine a g a r . Carbon uti lization: _D -Gluco s e , 4 -a rabino se, _D -xy lo s e, ,D -f r u c t o s e a n d raffinose a r e utilized f o r growth. N o growth o r only t r a c e of growth on _D-mannitol and r h a m n o s e . Utilization of s u c r o s e and--inositoL i s doubtful.
BIBLIOGRAPHY Ammann, A . , D . Cottlieb, T . D. B r o c k , H. E. C a r t e r and G. B . Whitfield. 1955. Filipin, a n antibiotic effective against fungi. Phytopathol. 45: 559-563. Anderson, H. W-.and D. Gottlieb. 1952. P l a n t d i s e a s e c o n t r o l with antibiotics. Econ. Bot. 6:294-308. Anderson, Lucia E., J . E h r l i c h , Sung Huang Sun a n d P. R. B u r k h o l d e r . 1956. S t r a i n s of S t r e p t o m y c e s , the s o u r c e s of a z a s e r i n e , elaiomycin, g r i s e o v i r i d i n , a n d v i r i d o g r i s e i n . Antibiotics a n d C h e m o t h e r s p y 6: 100-115. A r z , T . , T . Nakada and M. Suzuki. 1957. Production of v i o m y c i n Like s u b s t a n c e by a S t r e p t o m y c e s . Antibiotics and Chemotherapy 7: 435-442. Artamonova, 0. I. a n d N. A. Krasil'nikov. 1960. A c t i n o m y c e t e s of the violet group. (Aktinomitsety fioletovoy grupp). Biology of s p e c i a l g r o u p s of a c t i n o m y c e t e s - p r o d u c e r s of a n t i b i o t i c s (in R u s s i a n ) . T r u d . I n s t . Mikrobiol., Akad. Nauk, S.S.S.R. S: 274-337 (Engl. t r a n s l . p . 274-330). Backus, E . J . , H. D. T r e s n e r a n d T. H. Campbell. 1957. The nucleocidin a n d alazopeptin producing o r g a n i s m s : two new s p e c i e s of S t r e p t o m y c e s . Antibiotics a n d Chemotherapy 2: 532-541. B a l d a c c i , E. 1958. Development i n the c l a s s i f i c a t i o n of a c t i n o m y c e t e s . G i o r n . Microbiol. 5: 10-27. , G. F a r i n a a n d R. Locci. 1966. Emendation of the genus S t r e p t o v e r t i c i l l i u m B a l d a c c i (1958) a n d r e v i s i o n of s o m e s p e c i e s . 153-171. G i o r n . Microbiol. B a r r , F . S. a n d P. E . G a r m a n . 1956. S t r e p t o m y c e s kentuckensis, a new s p e c i e s , the p r o d u c e r of r a i s n o m y c i n . Antibiotics and c h e m o t h e r a p y 5: 286-289. Benedict. R. C . , W. Dvonch, Odette Shotwell, T. C . P r i d h a m a n d L. A. L i n d e n f e l s e r . 1952. Cinnamycin, a n antibiotic f r o m S t r e p t o m y c e s cinnamoneus nov. sp. Antibiotics and Chemotherapy 2 : 591- 594. , 0. L. Shotwelt, T. C . P r i d h a m , L. A. Lindenfelser and W. C. Haynes. 1954. The production of the neomycin complex by Streptomyces albogriseolus , 3 . Antibiotics and Chemotherapy 4: 653-656.
5:
x.
SYSTEMATIC
BACTERIOLOGY
38 3
B e r g e r , J . , L. M. Jampolsky, M. W. Goldberg. 1949. Borrelidin, a new antibiotic with anti-borrelia activity and penicillin enhancement p r o p e r t i e s . Arch. Biochem. 22: 476-478. B e s s e l l , Christine J . , W . K . Anslow, Aileen M. Mortimer, D. L. F l e t c h e r and A. Rhodes. 1961. Antibiotic production. U . S . P a t . 3,013, 947 Dec. 19. U.S. P a t . Office. 554-56t. B r u m p t , E . 1906. L e s Myc6 t o m e s . Arch. P a r a s i t o l . 2: Conn, H.J. 1917a. Soil flora studies. V. Actinomycetes in soil. N.Y. Agr. Exp. Sta. Tech. Bull. 60: 1-16. , 1917b. Soil flora studies. V. Actinomycetes i n soil. N. Y . Agr. Exp. Sta., Annual Rept. 36: 157-170. Davisson, J. W . and A. C. Finlay. 1959. Catenulin and i t s production. U.S. P a t . 2, 895,876 July 21. U.S. P a t . Office. Duch6, J . 1934. L e s actinomyces du groupe albus. Encycl. Mycol., VI. P a u l LeChevalier and Sons, P a r i s . 1-375. Duggar, B . M. 1948. Aureomycin: a product of the continuing s e a r c h 177-181. f o r new antibiotics. Ann. N.Y. Acad. Sci. 3: 1949. Aureomycin and preparation of s a m e . U.S. P a t . 2,482,055. Feb. 11. U . S . P a t . Office. Dvonch, W . , Odette Shotwell, R. C. Benedict, T.G. P r i d h a m and L.A. Lindenfelser. 1954. Antibiotics and Chemotherapy 4: 1135-1142. Erikson, D. 1935. The pathogenic a e r o b i c o r g a n i s m s of the Actinomyces group. Med. Res. Council ( B r i t . ) Spec. Rep., S e r . 203: 5-61. F a l c z o d e Morais, J . O . and Maria Dblia Maia. 1961. Uma contribui5;o a o estudo t a x o n h i c o do gen8ro Streptomyces - uma tentativa d e simplificafzo. Rev. Inst. Antibiot. 2 : 33-60. , Maria Ddlia Maia and Maria E. Souto Maior Genn. 1958. Sabre uma variedade d e Streptomyces comum nos s o l o s do Brasil: Streptomyces venezuelae v a r . r o s e o s p o r i nov. v a r . Rev. Inst. Antbiot. 99-106. Finlay, A. C., F.A. Hochstein, B .A. Sobin and F . X . Murphy. 1951. Netropsin, a new antibiotic produced by a Streptomyces. J. A m e r . Chem. SOC. 73: 341-343. and B .A. Sobin. 1952. Netropsin and p r o c e s s f o r i t s p r o duction. U . S . P a t . 2,586,762 F e b . 19. U . S . P a t . Office. F r i e d , J., D. P e r l m a n , A . F . Langlykke and E . 0 Titus. 1958. 16 alpha-hydroxylation steriods. U . S . P a t . 2, 855, 343 Oct. 7, U . S . P a t . Office. F r o m m e r , W. 1959. Zur Systematik d e r Actinemycin bilden Streptomyceten. Arch. Mikrobiol. 32: 197-206. G a r d n e r , A. D. and E . Chain. 1942. Proactinomycin: a Ilbacteriostatic" produced by a species of Proactinomyces. B r i t . J. Exp. Pathol. 23: 123-127. Gauze, G. F., T. P. Preobrazhenskaya, E.S. Kudrina, N . 0. Blinov, I.D. Rjabova and M.A. Sveshnikova. 1957. P r o b l e m of classification of actinomycetes -antagonists (in Russian). Government Publishing House of Medical Literature, Medgiz, Moscow. 1-398. Gottlieb, D. 1968. Communication f r o m G . F . Gauze. Int. J . Syst. Bacteriol. E: 20. ,P.K. Bhattacharyya, H. E. C a r t e r and H. W. Anderson. 1951, Endomycin, a new antibiotic. Phytopathol. %:393-400. , and H. E. C a r t e r . 1956. Endomycin and p r o c e s s f o r i t s production. U.S. P a t . 2, 746,902 May 22. U . S . P a t . Office.
r:
38 4
, a n d E. Shirting. 1967. Cooperative description of type c u l t u r e s of Streptomyces. I. The International Streptomyces P r o j e c t . Intern. J. Syst. Bacteriol. 315-322. Cougerot, H., P. B l u m and J. Duch6. 1934. Kerionactinomycosique du a Nocardia bovis. Arch. Dermato-Syphiligraphiques de la Clinique d e 1'Hop. : 376. CUSSOW, H. T. 1914. The systematic position of the o r g a n i s m of the common potato scab. Science 2: 431-433. Hamada, S . 1958. A study of a new antitumor substance, cellostatin. 1. On the isolation a n d some p r o p e r t i e s of cellostatin. Tohoku J . Exp. Med. 67:173-179. Hata, T., N . Ohki and T. Higuchi. 1952a. Studies on the antibiotic substance llluteomycinl'. On the s t r a i n s and the cultural conditions (in Japanese, English summary). J . Antibiot. 5: 529-534. , N. Ohki and T. Higuchi. 1952b. Studies on the antibiotic substance llluteomycinll. On the s t r a i n and the fermentation. Kitasato Arch. Exp. Med. 24: 67- 77 (447-45?). , N. Ohki and T. Higuchi. 1 9 5 2 ~ . Taxonomic studies on Streptomyces. IV. Correlation between the classification by utilization of carbon compounds and s p e c t r a on the a g a r plate and a n t i streptomycete s p e c t r a . Kitasato Arch. Exp. Med. 25: 188-189. HUtter, R. 1967. Systematik d e r Streptomyceten. S. K a r g e r , B a s e l . 382p. I m a m u r a , A . , M. Hori, K. Nakazawa, M. Shibata, S. Tatsuoka and A. Miyake. 1956. A new s p e c i e s of Streptomyces producing dihydrostreptomycin. P r o c . J a p . Acad. , Tokyo. 32: 648-653. Isono, K. , K. Anzai and S. Suzuki. 1958. T u b e r m y c i . A and B, new antibiotics, I. J. Antibiot., Tokyo S e r . A, L:264-266. Johnstone, D.B. and S.A. Waksman. 1947. Streptomycin II, a n a n t i biotic substance produced by a new species of Streptomyces. P r o c . SOC.Exp. Biol. and Med. 65: 294-295. , and S. A. Waksman. 1948. The production of Streptomycin by Streptomyces bikiniensis. J. B a c t e r i o l . 55: 317-326. Kawato, M. and R. Shinobu. 1959. On Streptomyces h e r b a r i c o l o r nov. sp. Supplement: a simple technique f o r the microscopical observation. Mem. Osaka Univ. Lib. A r t s and Educ. , B . Natur. Sci. 8: 114-119. Kondo, ST, T. Miyakawa, H. Yumoto, M. Sezaki, M. Shimura, K. Sato and T . Hara. 1962. Isolation and characterization of a new a n t i biotic, Bandamycin B . J. Antibiot., Tokyo S e r . A, &: 161-167. , J.M.J. Sakamoto and H. Yumoto. 1961. J. Antibiot., : 365-366. Tokyo S e r . A, E Korenyako, A.I., N.A. Krasil'nikov and N. I. Nikitina. 1960. Antinomyces levoris, Biology of special groups of actinomycetes producer3 of antibiotics (in Russian). Trud. Inst. Mikrobiol. , Akad. Nauk. S.S. S ,R. 8:116-132. (Engl. t r a n s l . p. 111-127). , N.A. Krasil'nikov, N.I. Nikitina and A.I. Sokolova. 1960. Actinomycetes of the fluorescent group. Biology of special groups of actinomycetes p r o d u c e r s of antibiotics (in Russian). Trud. Inst. Mikrobiol., Akad. Nauk. S.S.S,R 8: 133-159 (Engl. t r a n s l . p. 128-156). Krainsky, A. 1914. Die Aktinomyceten und i h e r e n Bedeutung i n d e r 649-688. Natur. Zbl. Bakt. Abt. 2 ,
c:
s:
SYSTEMATIC
BACTERIOLOGY
38 5
K r a s i l ' n i k o v , N.A. 1941. Guide t o t h e a c t i n o m y c e t e s (in R u s s i a n ) . Akad. Nauk. S . S . S , R . , Moscow. 1-158. , 1949. Guide t o t h e b a c t e r i a a n d a c t i n o m y c e t e s (in R u s s i a n ) . Akad. Nauk. S . S . S . R . , Moscow. 1-830. , 1955. T r a n s . Int. Symp. on Antibiotics, W a r s a w : 12-18. , 1957. La c l a s s i f i c a t i o n d e s a c t i n o m y c e t e s p r o d u c t e u r s d'antibiotiques. Ann. I n s t . P a s t e u r 2: 597-604. , a n d N.S. A g r e . 1960. A c t i n o m y c e t e s of the white-blue group. Biology of s p e c i a l g r o u p s of a c t i n o m y c e t e - p r o d u c e r s of a n t i b i o t i c s . (In R u s s i a n ) . T r u d . I n s t . Mikrobiol. Akad. Nauk. S. S. S.. L,, 8: 254-274. Kuchaeva, A.G., N.A. Krasil'nikov, G.K. S k r y a b i n a n d S . D . Taptykova. 1960. A c t i n o m y c e t e s of the olivochromogenous g r o u p s . (Burozelenoy grupp). Biology of s p e c i a l g r o u p s of a c t i n o m M S W R 3 E H y c e t e s - p r o d u c e r s of a n t i b i o t i c s (in R u s s i a n ) . T r u d . I n s t . Mikrobiol., Akad. Nauk S.S.S. R, S: 226-253 (Engl. t r a n s l . p. 222-251). , N.A. Krasil'nikov, S . D . Taptikova a n d R. Z . G e s h e v a . 1962. On t h e taxonomy of a c t i n o m y c e t e s of the Lavendulae g r o u p (in R u s s i a n , E n g l i s h s u m m a r y ) . Bull. I n s t . Microbiol. Sofia 13: 103-124. Kudrina, E . S. 1957. C h a r a c t e r i s t i c s of a n t a g o n i s t i c a c t i n o m y c e t e s of the Helvolus s e r i e s (in R u s s i a n ) , p. 77-100. Z G a u z e , G . F . , T. P. P r e o b r a z h e n s k a y a , E. S. Kudrina, N. 0. B linov, E. D. Ryab ova a n d M.A. Sveshnikova. P r o b l e m s of c l a s s i f i a c t i o n of a c t i n o m y c e t e s a n t a g o n i s t s . G o v e r n m e n t P u b l i s h i n g House of Medical L i t e r a t u r e , Medzig.. Moscow. K u t z n e r , H. J . a n d S.A. Waksman. 1959. S t r e p t o m y c e s c o e l i c o l o r Mtlller a n d S t r e p t o m y c e s v i o l a c e o r u b e r W a k s m a n a n d C u r t i s , two 528-538. d i s t i n c t l y d i f f e r e n t o r g a n i s m s . J. B a c t e r i o l . 2: Leben, C., G . J. S t e s s e l a n d G . W. Keitt. 1952. Helixin, a n a n t i biotic a c t i v e a g a i n s t c e r t a i n fungi a n d b a c t e r i a . Mycologia 159-168. L i e s k e , R. 1921. Morphologic und biologic d e r S t r a h l e n p i l z e . B o r n t r a g e r , Leipzig McGuire, J. M. 1954. I m p r o v e m e n t s i n o r r e l a t i n g to T e r r a m y c i n . B r i t i s h P a t . Specification 7L3, 795 Aug. 18. P a t . Office, London. Maeda, K., H. Kosaka, Y . O k a m i a n d H. Umezawa. 1953. A new antibiotic, p y r i d o m y c i n . J. Antibiot., Tokyo S e r . A, 5: 140. Margalith, P. a n d G. B e r e t t a . 1960. A new antibiotic producing S t r e p t o m y c e s : E r . b e l l u s nov. SE. Mycopathol. e t Mycol. Appl. 12 : 189-195. , C. B e r e t t a a n d M . T . T i m b a l . 1959. Matamycin, a new antibiotic. I. Biological s t u d i e s . Antibiotics a n d C h e m o t h e r a p y 9: 71-75. M i l l a r d , W.A. a n d S. B u r r . 1926. A study of twenty-four s t r a i n s of A c t i n o m y c e s a n d t h e i r r e l a t i o n to t y p e s of c o m m o n s c a b of potato. 580-644. Ann. Appl. B i o l . g: Mtlller, R. 1908. E i n e Diphtheridee und e i n e S t r e p t o t h r i x m i t g l e i c h e m blauen F a r b s t o f f sowie U n t e r s u c h u n g e n tlber S t r e p t o t h r i x a r t e n im a l l g e m e i n e n Zbl. B a k t . , Abt. I. O r i g . 46: 195-212.
s:
38 6
Nagatsu, J . , K. A n s a i , K. Ohkuma a n d S . Suzuki. 1963. S t u d i e s on a new a n t i b i o t i c , t u b e r i n . IV. T a x o n o m i c s t u d i e s on t h e t u b e r i n p r o ducing o r g a n i s m s , S t r e p t o m y c e s a m a k u s a e n s i s . J . Antibiot. , Tokyo S e r . A, 207-210. N a k a m u r a , G . 1961. S t u d i e s on a n t i b i o t i c a c t i n o m y c e t e s . I. On S t r e p t o m y c e s p r o d u c i n g a new a n t i b i o t i c t u b e r m y c i n . J. Antibiot., Tokyo S e r . A,. E: 86-89. Nakazawa, K. 1964. S t u d i e s on s t r e p t o m y c e t e s . M i c r o b i o l o g i c a l c h a r a c t e r i s t i c s of S t r e p t o m y c e s no. 5866. Annu. Rep. T a k e d a Res. L a b . 23: 69-70. , e t a t . 1954. Candimycin. J a p . P a t . 2125 F e b . 4. , a n d S . F u j i i . 1957. S t u d i e s on s t r e p t o m y c e t e s . On S t r e p t o m y c e s s i n d e n e n s i s nov. sp. Annu. Rep. Takeda R e s . L a b . 16: 109-110. , S . F u j i i , M. Inoue, H. Hitomi, A. Miyake a n d I. Kaneko. 1954. On a new a n t i - t u b e r c u l o u s s u b s t a n c e p r o d u c e d by a S t r e p t o - m y c e s (in J a p a n e s e ) . J. Antibiot., Tokyo Ser. B, 168. , a n d M. Shibata. 1956. S t r e p t o m y c e s humidus nov. s p . , p . 648. k I m a m u r a , A . , M. Hori, K. Nakazawa, M, Shibata, S. T a t s u o k a a n d A. Miyake. 1956. A new s p e c i e s of S t r e p t o m y c e s p r o d u c i n g d i h y d r o s t r e p t o m y c i n . P r o c . J a p . A c a d . , Tokyo, 32: 648-653. , M. Shibata, M. Honjo, Y . H a m a d a , J. Kaneko a n d Y . Tokui. 1956. Candimycin. J a p . P a t . 8148 Nov. LO. , M. Shibata, K . T a n a b e , A. Miyake, K . Mizuno a n d T. Takewaka. 1957. S t u d i e s on s t r e p t o m y c e t e s . On S t r e p t o m y c e s l i l a c i n u s nov. s p . a n d i t s a n t i b i o t i c c l a d o m y c i n . Annu. Rep. T a k e d a R e s . Lab. 111-115. , M. Shibata, K. T a n a b e a n d H. Y a m a m o t o . 1958. S t u d i e s on s t r e p t o m y c e t e s . S t r e p t o m y c e s h u m i d u s nov. sp. p r o d u c i n g d i h y d r o s t r e p t o m y c i n . J . A g r . C h e m . SOC.J a p a n 32: 321-324. , M. Shibata, K. T a n a b e a n d H . Y a m a m o t o . 1960. P r o d u c tion of d i h y d r o s t r e p t o m y c i n . U.S. P a t . 2, 931, 756 A p r . 5. U.S. P a t . Office. (Also, U . S . P a t . 2, 950, 277 Aug. 23). , Y T a n a b e , M. Shibata, A. Miyake a n d T. Takewaka. 1955. S t u d i e s on s t r e p t o m y c e t e s . C l a d o m y c i n a new a n t i b i o t i c p r o d u c e d by S t r e p t o m y c e s l i l a c i n u s nov. s p . (in J a p a n e s e ) . J . Antibiot. S e r . B, 9: 81. Niida, T . J a p a n e s e P a t . 425900. A b s t r . i n J a p a n Offic. G a z . 38-26948. Nikitina, N.N. 1958. A c t i n o m y c e t e s of t h e G l o b i s p o r u s g r o u p . A b s t r . of T h e s i s . Mikrobiologia 27: 141. A. I . K o r e n y a k o a n d N . A . K r a s i l ' n i k o v . 1960. A c t i n o m y c e t e s t r a i n s of t h e s p e c i e s A c t i n o m y c e s s t r e p t o m y c i n i K r a s s . Biology of special g r o u p s of a c t i n o m y c e t e s - p r o d u c e r s of a n t i b i o t i c s (in R u s s i a n ) . T r u d . I n s t . Mikrobiol., Akad, Nauk. S . S . S . R . 8: 85-103 (Engl. t r a n s l . p . 80-98). N i s h i m u r a , H . , T. K i m u r a , K. T a w a r a , K. S a s a k i , K. N a k a j i m a , N. Shimaoka, S. Okamodo, M. S h i m o h i r a a n d J. I s o n o . 1967. A b u r a m y c i n , a new a n t i b i o t i c . J. A n t i b i o t i c s , Tokyo S e r . A,
1 6 :
L:
1 6 :
10: -
205-211.
387
Okami, Y . , K. Maeda a n d H. Umezawa. 1954. Studies on antibiotic a c t i n o m y c e t e s , VIII. On S t r e p t o m y c e s a l b i d o f u s c u s O k a m i g Umezawa, nov. s p . producing a new a n t i b i o t i c p y r i d o m y c i n . J . Antibiot., Tokyo S e r . A, 2: 55-56. , a n d H. U m e z a w a . 1955. & O k a m i , U t a h a r a , Oyagi, N a k a m u r a , Umezawa, Yanagisawa a n d T u n e m a t s u . The s c r e e n i n g of a n t i t o x o p l a s m i c s u b s t a n c e p r o d u c e d by S t r e p t o m y c e t e a n d a n t i - t o x o p l a s m i c s u b s t a n c e No. 534. J . Antibiot., Tokyo S e r . A, S: 126-131. , a n d H . Umezawa. 1959. N u r a s e e t a l . Identification of a n antibiotic p r d u c e d by 2. galbus, n . sp. with s t r e p t o m y c i n . J. Antibiotics, Tokyo S e r . A, 126-132. , R . U t a h a r a , S . N a k a m u r a a n d H. U m e z a w a . 1954. S t u d i e s on antibiotic a c t i n o m y c e t e s , IX. On S t r e p t o m y c e s producing a new antifungal s u b s t a n c e mediocidin a n d antifungal s u b s t a n c e s of fungicidin - r i m o c i d i n - c h r o m i n group, e u r o c i d i n g r o u p a n d t r i c h o m y c i n a s c o s i n - c a n d i c i d i n g r o u p . J. Antibiot., Tokyo S e r . A, 1:98-100. Oswald, Elizabeth, J . , R. J . Reedy and W.A. Randall. 1956. An a n t i fungal agent, 1968, p r o d u c e d by a new S t r e p t o m y c e s s p e c i e s , p . 236239. & A n t i b i o t i c s Annual 1955-L956. Medical Encyclopedia I n c . , New Y o r k . P e r l m a n , D . , J. F r i e d , E . O . T i t u s a n d A . F . Langlykke. 1955. 16 alpha-hydroxyprogesterone, 16 alpha-hydroxydihydroprogesterone a n d e s t e r s t h e r e o f . U.S. P a t . 2, 709, 705 May 31, U.S. P a t . Office. P i n n e r t - S i n d i c o , Sylvie. 1954. Une nouvelle e s p & c e d e S t r e p t o m y c e s p r o d u c t r i c e d'antibiotiques: S t r e p t o m y c e s a m b o f a c i e n s n. sp. 702-708. c a r a c t g r e s c u l t u r a u x . Ann. I n s t . P a s t e u r 87: P i t t e n g e r , R.C. a n d D. Gottlieb. 1961. p . 258. Waksman, S.A. 1961. The A c t i n o m y c e t e s . V o l . 2. C l a s s i f i c a t i o n , identification a n d d e s c r i p t i o n of g e n e r a a n d s p e c i e s . The WilLiams and Wilkins Co., B a l t i m o r e . L - 363. P r e o b r a z h e n s k a y a , T. P. 1957a. C h a r a c t e r i s t i c s of a n t a g o n i s t i c a c t i n o m y c e t e s of the F r a d i a e s e r i e s (in R u s s i a n ) , p. 51-57. & G a u z e , G . F . , T . P . P r e o b r a z h e n s k a y a , E.S. Kudrina, N.O. B l i n w , I.D. Ryabova a n d M.A. Sveshnikova. P r o b l e m s of c l a s s i f i c a t i o n of a c t i n o m y c e t e s - a n t a g o n i s t s . G o v e r n m e n t Publishing House of Medical L i t e r a t u r e , Medgiz, Moscow. , 1957b. C h a r a c t e r i s t i c s of a n t a g o n i s t i c a c t i n o m y c e t e s of t h e C o e r u l e s c e n s s e r i e s (in R u s s i a n ) , p. 119-130. & G a u z e , G . F . , T . P. P r e o b r a z h e n s k a y a , E.S. Kudrina, N. 0. Blinov, I.D. Ryabova a n d M.A. Sveshnikova. P r o b l e m s of c l a s s i f i c a t i o n of a n t i n o m y c e t e s a n t a g o n i s t s . G o v e r n m e n t P u b l i s h i n g House of Medical L i t e r a t u r e , Medgiz, Moscow. , 1 9 5 7 ~ . C h a r a c t e r i s t i c s of a n t a g o n i s t i c a c t i n o m y c e t e s of the R u b e r s e r i e s (in R u s s i a n ) , p. 72-76. & G a u z e , G . F . , T . P . P r e o b r a z h e n s k a y a , E . S . Kudrina, N. 0. Blinov, I. D. Ryabova a n d M.A. Sveshnikova. P r o b l e m s of c l a s s i f i c a t i o n of a c t i n o m y c e t e s a n t a g o n i s t s . G o v e r n m e n t Publishing House of Medical L i t e r a t u r e , Medgiz, Moscow. , N. 0. Blinov a n d 1. D. Ryabova. 1957. C h a r a c t e r i s t i c s of a n t a g o n i s t i c a c t i n o m y c e t e s of the G r i s e u s s e r i e s (in Russian), p . 131145. & G a u z e , G . F . , T , P . P r e o b r a z h e n s k a y a , E.S. Kudrina, N . O . Blinov, I. D. Ryabova a n d M . A . Sveshnikova. P r o b l e m s of c l a s s i f i cation of a c t i n o m y c e t e s - a n t a g o n i s t s . G o v e r n m e n t P u b l i s h i n g House of Medical L i t e r a t u r e , Medgiz, Moscow.
c:
388
INTERNATIONAL
JOURNAL
, I.D. Ryabova and N.O. Blinov. 1957. C h a r a c t e r i s t i c s of antagonistic actinomycetes of the Chromogenes s e r i e s (in Russian) p. 169-177. &Gauze, G . F . , T . P . Preobrazhenskaya, E. S. Kudrina, N . O . Btinov, I . D . Ryabova and M.A. Sveshnikova. P r o b l e m s of classification of actinomycetes -antagonists. Government Publishing House of Medical L i t e r a t u r e , Madgiz, Moscow. , a n d M.A. Sveshnikova. 1957. C h a r a c t e r i s t i c s of antagonistic actinomycetes of the Lavendulae- RoseQs s e r i e s (in Russian), p. 31-50. &Gauze, C . F. , T. P. Preobrazhenskaya, E. S . Kudrina, N.O. Blinov, I.D. Ryabova and M.A. Svenshnikova. P r o b l e m s of classification of actinomycetes -antagonists. Government Publishing House of Medical L i t e r a t u r e , Medgiz, Moscow. P r i d h a m , T.G., C. W. Hessettine and R.G. Benedict. 1958. A guide f o r the classification of streptomycetes according to selected groups: placement of s t r a i n s in morphological sections. Appl. Microbiol. 6:52 - 79. , Odette Shotwell, F . H . Stodola, L.A. Lindenfelser, R.G. Benedict and R. W. Jackson. 1956. Antibiotics against plant d i s e a s e . II. Effective a g e n t s produced by Streptomyces cinnamomeus f o r m a azacoluta f . nov. Phytopathology 46: 575-581. Reynolds, D. M. and S.A. Waksman. 1948. Grisein, a n antibiotic p r o duced by c e r t a i n s t r a i n s of Streptomyces g r i s e u s . J. Bacteriol. 55: 739-752. Ryabova, I. D. and T. P. Preobrazhenskaya. 1957. C h a r a c t e r i s t i c s of antagonistic actinomycetes of the Violaceus s e r i e s (in Russian), p. 178-197. &Gauze, C . F . , T. P. Preobrazhenskaya, E.S. Kudrina, N . O . Blinov, I.D. Ryabova and M.A. Sveshnikova. P r o b l e m s of classification of actinomycetes-antagonists. Government Publishing House of Medicat L i t e r a t u r e , Medgiz, Moscow. Sakamoto, J e a n Marie J . , S. Kondo, H. Yumoto and M. A r i s h i m a . 1962. Bundlins A and B , two antibiotics produced by Streptomyces 98-101. griseofuscus nov. sp. J. Antibiot., Tokyo S e r . A, 2: Sawazaki, T . , S. Susuki, G . Nakamura, M. Kawasaki, S . Yamashita, K. Isono, K. Anza;, Y . Serizawa and Y . Sekiyama. 1955. Streptomycin production by a new strain, Streptomyces m a s h u e n s i s J. Antibiot., Tokyo S e r . A, 2: 44-47. Shibata, M. , M. Honj6, Y . Tokui and K. Nakazawa. 1954. On a new antifungal and anti-yeast substance, Candimycin, produced by a s t r e p t o m y c e s . (In Japanese). J. Antibiot., Tokyo S e r . B , 168. , K. Nakazawa, A. Miyake, M. Inoue and A. Okabori. 1957. Studies on streptomycetes. Croceomycin, a new antituberculous 32-37. substance. Annu. Rep. Takeda Res. Lab. Shimo, M., T. Shiga, T. Tomosugi and I . Kamoi. 1959. Studies on taitomycin, a new antibiotics produced by Streptomyces, sp. no. 772 afghaniensis). I. Studies on the s t r a i n and production of taitomycin. J. Antibiot., Tokyo S e r . A, 2:1-6. Shinobu, R. 1956. T h r e e new s p e c i e s of Streptomyces forming w h i r l s . Mem. Osaka Univ. Lib. A r t s and Educ., B . Nat. Sci., No. 2: 84-93. , 1957. Two new s p e c i e s of Streptomyces Mem. Osaka Univ. Lib. A r t s and Educ., B. Natur. Sci. 5: 63-73. , and M. Kawato. 1960a. On Streptcmyces aerocolonigenes nov. s p . , forming the secondary colonies on the a e r i a l mycetia. Bot. Mag., Tokyo 73: 212-216.
L:
1 6 :
(c.
SYSTEMATIC
BACTERIOLOGY
389
, and M. Kawato. 1960b. On Streptomyces indigoferus nov. sp. producing blue to g r e e n soluble pigment on some synthetic media. Mem. Osaka Univ. Lib. A r t s and Educ., B . Natur. S c i . 9: 49-53. Shirling, E . B . and D. Gottlieb. 1966. Methods f o r characterization of Streptomyces species. Intern. J . of Systemat. Bacteriol. 313 - 340. , and D. Gottlieb. 1968. Cooperative description of type cultures of Streptomyces. II. Species descriptions f r o m f i r s t study. Intern. J. Syst. Bacteriol. g: 69-189. Sobin, B.A., A.C. Finlay and J.H. Kane. 1950. T e r r a m y c i n and i t s production. U.S. P a t . 2,516,080 July 18. U.S. P a t . Office. Stark, W.M., C.E. Higgens, R.N. Wolfe, M.M. &ehn and J.M. McGuire. 1962. Capreomycin, a new antimycobacterial agent produced by Streptomyces capreolus sp. n. Antimicrobial Agents and Chemotherapy 1962: 596-606. Sugai, T. 1956. New antibiotics 229 and 229 B of c o l o r l e s s , watersoluble nature (in Japanese). J . Antibiot., Tokyo Ser. B, ?: 170 - 179. Sugawara, R. and M. Onuma. 1957. Melanomycin, a new antitumor substance f r o m Streptomyces. II. Description of the strain. J. Antibiot., Tokyo S e r . A, Lo: 138-142. Suzuki, S., G. Nakamura, K. Okuma and Y . Tomiyama. 1958. 81-83. Cellocidin, a new antibiotic. J . Antibiot., Tokyo S e r . A, Sveshnikova, M.A, C h a r a c t e r i s t i c s of antagonistic actinomycetes of the Nigrescens s e r i e s (in Russian), p. 146-148. &Gauze, G . F . , T. P. Preobrazhenskaya, E.S. Kudrina, N.O. Blinov, I.D. Ryabova and M.A. Sveshnikova. P r o b l e m s of classification of actinomycetesantagonists. Government Publishing House of Medical Literature, Medgiz, Moscow. Szab6, I., Maria Marton, Ilona Buti and G. P d r t a i . 1963. Actinomyces finlayi n - sp. Acta Microbiol. Acad. Sci. Hungaricae 10: 207-214. Thaxter, R. 1892. Potato scab. Conn. Agri. Exp. Sta. Annu. Rep. f o r 1891: 153-160. Trejo, W. 1961. Streptomyces nodosus. p. 250. & Waksman, S.A. 1961. The Actinomycetes. Vol. 2. Classification, identification and description of genera and species. The Williams and Wilkins Co. , Baltimore. 1-363. , and R.E. Bennett. 1963. Streptomyces nodosus sp. n., the amphotericin-producing organism. J. Bacteriol. 85: 436-439. T r e s n e r , H.D., M.C. Davies and E. J. Backus. 1961. Electron microscopy of streptomyces s p o r e morphology and i t s r o l e in species differentiation. J. Bacteriol. 2:70-80. Thirumalachar, M.J. and V . V . Bhatt. 1960. Some Streptomyces species producing oxytetracycline. Hindustan Antiobiot. Bull. 3: 61-63. Umezawa, H. and Y . Okami. 1957. Changing the name of pyridomycin producing streptomycete. J. Antibiot. , Tokyo Ser. A, Lo: 172. Upjohn Company. 1954. The antibiotic endomycin and p r o c e s s f o r i t s production. B r i t . Pat. Spec. 705,622 March 17. Patent Office, London. Waksman, S.A. 1919. Cultural studies of species of Actinomyces. Soil Sci. S: 71-215.
1 6 :
1 1 :
39 0
INTERNATIONAL
JOURNAL
, 1953. Waksman and LechevaLier. Guide to the classification and identification of the actinomycetes and their antibiotics. The Williams and Wilkine Co. , Baltimore. 1-162. 1957. Family Actinomycetaceae Buchanan and family Streptomycetaceae Waksman and Henrici Breed, Murray and Smith, Bergey's Manual of Determinative Bacteriology, 7th ed. , 744 825. 1959. Strain specificity and production of antibiotic substances. X. Characteriaztion and classification of species within the Streptomyces g r i s e u s group. P r o c . Nat. Acad. Sci. U.S. 2: 1043- 1046. 1961. The actinomycetes. Vol. 2. Classification, identification and descriptions of genera and species. The Wiltiams and Wilkins Company, Baltimore. 1-363. , and R.E. Curtis. 1916. The actinomyces of the soil. Soil Sci. I= 99-134. , and A. T. Henrici. 1948. Family Actinomycetaceae Buchanan and Fami ly Streptomycetaceae Waksman and Henrici p. 892-980. &Breed, Murray and Hitchens, Bergey's Manual of Determinative Bacteriology, 6th ed., The Williams ane Wilkins Go., Baltimore. , Etiz. S. Horning, M. Welsch and H. B. Woodruff. 1942. Distribution of antagonistic actinomycetes in nature. Soil Sci. 54: 281 290. , and H.B. Woodruff. 1941. Actinomyces antibioticus, a new soil organism antagonietic to pathogenic and non-pathogenic bacteria. J. Bacteriol. 42: 231-249. . , D.A. Harris, Mary Ann Regan, Myrle Ruger and H.B. Wallick, H Woodruff. 1956. Discovery and antimicrobial p r o p e r t i e s of cathomycin, a new antibiotic produced by Streptomyces spheroides n. sp. , p. 909-916. &Antibiotics annual 1955-1956. Medical Encyclopedia Inc., New York. Wooldridge, W.E. 1957. A new fungal antibiotic NC 1968. German Pat. Not. (DAS) 1000966 Jan. 17. 1960a. Antifungal antibiotic designated NC 1968. British Pat. 828, 792, Feb. 24. L960b. Antifungal antibiotic and 5. coelicolor v a r . aminophilus. U.S. P a t . 2,956,925 Oct. 18. U.S. P a t . Office. Yamaguchi, T. and Y. Saburi. 1955. Studies on the anti-trichomonal actinomycetee and their classification. J. Cen. Appl. Microbiol.
.
.
1:
201-235.
SYSTEMATIC BACTERIOLOGY SOURCES OF ILLUSTRATIONS 1.

2.
3.
39 1
4.
5 .
6. 7. 8.
9. LO.
11.
12. 13. 14. 15.
16. 17.
18. 19.
20.
21.
22.
23.
24.
G.F. Gauze. Acad. Med. Sciences of USSR, Institute of New Antibiotic s, MOs c ow , USSR. ELio Baldacci, G . Farina and R. Locci. Instituto Di Patologie, Vegetale dell' Universita', V i a Celoria N. 2, Milano, Italia. H. P r a u s e r . Inst. Mikrobiol. Exptl. Therapie, Beuthenbergstr. 11 Jena, German Democratic Republic. E. J. Rdzok and T. J. Oliver. Abbott Laboratories, N. Chicago, Illinois,. U.S.A. Y . Ohara and H. Nonomura. Faculty of Engineering, Yamanashi Univ. , Kofu, Japan. J. Mathews. The Upjohn Co., Kalamazoo, Mich. 49001, U.S.A. A. Feltynowski and Wanda Wofnika. Panetwowy Zaktad Higieny, U1. Chocimska 24, Warszawa 12, Poland. T. C r o s s and Miss A. Maciver. Bradford Institute of Technology, Bradford 7, Yorkshire, England. Mr. Oates. Dept. of Bot., Leeds Univ., Leeds, Yorkshire, England. N.A. Krasil'nikov. Inst. Microbiol. , Acad. Sciences, MOSCOW, USSR. G.A. deVries. Certtaalbureau voor Schimmelcultures, Baarn, Nether lands. G.V. Mierlo and A. L. Houwink. Lab. f. Microbiol., Delft, Nether lands. (J. ElLiott) J. Spyvee. Boots P u r e Drug Co., Ltd., Antibiot. and F e r m . Div. , Nottingham, England. K. Nakazawa. Takeda Chem. Ind. , Ltd. , Juso-Higashiyodogawaku, Osaka, Japan. J. Fiala and Alena &ciEovd. Czech. Acad. of Science, Inst. of Mikrobiol., Prague 4, Budejovicka No. 1083, Czechos lovakia. W.A. Trejo. E.R. Squibb and Sons, New Brunswick, New J e r s e y , U.S.A. M. Mayama and K. Terada. Div. of Bacteriol., Shionogi Research Lab. , Shionogi and CO. , Ltd. , Fukushima-ku, Osaka, Japan. Y. Okami. Nat'l Inst. of Health of Japan, Shinagawa, Tokyo, Japan. G. Michaels. Dept. Microbiol. , Univ. Georgia, Athens, Georgia, U. S . A . Ina Radziminska. Hoffman-LaRoche, Nutley, New J e r s e y , U.S.A. V.A. Tsyganov. Res. Inst. of Antibiotics, Leningrad, L-20, USSR. T. R. Vernon. Dept. Scientific and Ind. Res. , P r i v a t e Bag, Auckland C1, New Zealand. S. T. Williams. Hartley Botanical Laboratories, Univ. of Liverpool, Liverpool, England. V. Pokornf and Alena kiciEovd. Czech. Acad. of Science, Inst. of Mikrobiol., Prague 4, Budejovicka No. 1083, Cze chos Lovakia
39 2
INTERNATIONAL
JOURNAL
25. 26.
27. 28.
29. 30.
31.
32.
33.
34. 35.
36.
37. 38. 39.
40.
Wanda Wognicka. Panstwowy Zakiad Higieny, U1. Chocimska 24, Warszawa 12. Poland. V.D. Kuznetsov. USSR R e s . Inst. for Antibiotics, MOSCOW, USSR. C a r o l S. Cassidy and K . E . Crook, Jr. B r i s t o l L a b s . , Inc. Syracuse, New York, U.S.A. S a r a Wold and Lucia Anderson. P a r k e , Davis and Go., Detroit, Michigan, U. S A . P. Lockwood. Dept. Textile Ind., B r a d f o r d Inst. of Technology, Bradford 7, Yorkshire, England. I. Szab6. Magyar Tudomdnyos AKaddmia Tatajtana h s Agrokdmiai Kutat6 IntBzeta, Budapest, II, Hungary. F. Mach. Inst. F u r Mikrobiologie, Griefswald. Ludwig-Jahn-Str., 15, G e r m a n Democratic Republic. J . Ludvik, Laborato; Elektronovd Mikroskopie. Mikrobiologickq K r ? ! , Budgjovicki 1083, Cziechoslavakia. 6 s l a v ESAV, P r a h a 4 T. J . Oliver. Abbott Laboratories, N. Chicago, Illinois, U.S.A. W. Hodgkiss. T o r r y Res. Sta., Aberdeen, Scotland. J. Rosillo and J. Spyvee. Boots P u r e Drug Go., Ltd., Antibiot. and F e r m . Div., Nottingham, England. J . Routien. C h a r l e s P f i z e r and Co., Inc., Groton, Conn.,
U.S.A.
C. E. Higgens, ELi Lilly and Go., Indianapolis, Indiana, U.S.A.
41.
M. Mayama and K. Tawara, Shionogi Res. Lab., Shionogi and Co., Ltd., Fukushima-ku, Osaka, Japan. J.O. Falcxo d e Morais, Inst. de Quimica da Univ. F e d . de Pe r na mbuc 0, Re c i f e , P e r na mbu c 0, B ra zi 1. A. Sokolov (and V. D. Kuznetsov). Lab. of Electronmicroscopy. Acad. Sci , USSR. N.M. McCLung. Dept. Bot. and Bacteriol., Univ. of South Florida, Tampa, Florida, U. S.A.

Cooperative Description of Type Cultures of STREPTOMYCES III. Additional Species Descriptions From First and Second Studies

Încărcat de

Informații document

Drepturi de autor

Formate disponibile

Partajați acest document

Partajați sau inserați document

Opțiuni de partajare

Vi se pare util acest document?

Este necorespunzător acest conținut?

Drepturi de autor:

Formate disponibile

Cooperative Description of Type Cultures of STREPTOMYCES III. Additional Species Descriptions From First and Second Studies

Încărcat de

Drepturi de autor:

Formate disponibile

INTERNATIONAL JOURNAL O F SYSTEMATIC BACTERIOLOGY October 1968 Vol. 18, No.

4 Copyright 1968, Iowa State University P r e s s

pp. 279- 392

R e v e r s e side of colony: No distinctive pigments (pale grayed yellow)

9 Leben, S t e s s e l and Keitt 1952, 159-168; Streptomyces antimycoticus

F i g u r e 24. S. a r g e n t e o l u s . Smooth s p o r e s ; e l e c t r o n m i c r o g r a p h f r o m 14 day c u l t u r e on o a t m e a l a g a r . lo

D- glucose may inaicate a r e q u i r e m e n t f o r a growth factor

P o o r growth on a l l carbon s o u r c e s including not p r e s e n t

is pH sensitive changing f r o m red-brown to yellow with addition of 0.05

Figure 80. S. gardneri. RF spore chains (X 600) on salts-starch agar,

F i g u r e 1 0 7 . 2. indigoferus. Smooth s p o r e s ; electron micrograph from 14 day culture on yeast-malt a g a r . 30

SYSTEMATIC of growth on L_-arabinose, raffinose.

BACTERIOLOGY _D-mannitol, rhamnose and

Figure 140. ,A. ni rescens agar, 21 days.

S i r a l spore chains (X 300) on salts-starch

_D-mannitol, _D-fructose and rhamnose a r e utilized for R e p o r t s v a r y on utilization of raffinose.

Streptomyces phaeoviridis Shinobu.

Description: Shinobu 1957, 67-

SYSTEMATIC BACTERIOLOGY SOURCES OF ILLUSTRATIONS 1.

12. 13. 14. 15.

S-ar putea să vă placă și