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

I N T E R N A T I O N A L J O U R N A L O F S Y S T E M A T I C B A C T E R I O L O G Y

Vol. 18, No. 4 October 1968 pp. 279- 392 Copyright 1968, Iowa State University P r e s s

COOPERATIVE DESCRIPTION O F TYPE CULTURES OF 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 1 8 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 - f o r t t o a s s e m b l e a n d r e d e s c r i b e a u t h e n t i c t y p e s t r a i n s o r a c c e p t a b l e n e o t y p e s t r a i n s f o r t h e n a m e d s p e c i e s i n t h e g e n e r a S t r e p t o m y c e s a n d S t r e p t o v e r t i c i l l i u m . P a r t I 1 o f 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 t h e r e p o r t t o i n c l u d e t y p e s t r a i n s f o r a n a d d i t i o n a l 1 0 0 S t r e p t o m y c e s o r S t r e p t o v e r t i c i l l i u m s p e c i e s . E a c h n e w d e s c r i p t i o n i s b a s e d u p o n d a t a s u b m i t t e d b y t h r e e c o - o p e r a t i n g l a b o r a t o r i e s , u s i n g t h e s t a n d a r d i z e d c r i t e r i a a n d m e t h o d s o f f i c i a l l y a d o p t e d f o r t h i s p u r p o s e . E m e n - d a t i o n s f o r a d d i t i o n a l s p e c i e s w i l l b e i n c l u d e d i n r e - p o r t s t o f o l l o w . T y p e . s p e c i m e n s f o r a l l r e d e s c r i b e d s p e c i e s a r e d e p o s i t e d i n t h e A m e r i c a n T y p e C u l t u r e 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 U S S R 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 J a p a n .

- - - - - - - - - - - -

Collaborators and authors of descriDtions:

Group A- 2*

Cross, T. and A. Maciver. Postgraduate School of Studies in Biological Sciences, Univ. of Bradford, Bradford 7 / Yorkshire, England.

Johanides, Vera and Tea Blazekovic. Laboratorij za industrijsku mikrobiologiju, Tehnoloxki fakultet, Pierottijeva 6, Zagreb, Jug0 s lavij a.

Trejo, W. E. R. Squibb and Sons, New Brunswick, New Jersey , U. S. A.

*: A letter-number combination is a s signed to each collaborating group. To conserve space, this code is used to indicate the author group for each emended description. The letter A indicates collaborating groups for the first 130 type s t ra ins distributed for study; B designates groups participating in the second study. ** Reprints of this art icle may be purchased f rom the International Micr.obiologica1 Fund, 22 1 Science Hall, Iowa State University, Ames, Iowa 50010 U. S. A. P r i ce for single copies: 562.00.

280 I N T E R N A T I O N A L J O U R N A L

Group A-3

Crook, Kenneth E., Jr. and Caro l S. Cassidy. Br i s to l L a b s . , Inc.

deVr ies , G. A. Centraalbureau voor Schimmelcultures, Baarn,

Krasil 'nikov, N. A. Inst. Microbiology. Academy of Sciences, Moscow

Syracuse, New York, U. S. A.

Netherlands.

B- 133, U . S. S. R.

Group A-8

Oliver, T. J. Abbott Labora tor ies , N . Chicago, Illinois, U. S. A. Spyvee, J. and J. Elliott, Boots P u r e Drug C o . , Ltd., Antibiot. and

Nakazawa, K. , M. Shibata, K. Yamamoto, E. Higashide, T . Iwasa and F e r m . Div., Nottingham, England.

T . Hasagawa. Takeda Chem. Ind., Ltd., Juso-Higashiyodogawaku, Osaka, Japan.

Group A-9

Nishimura, H., M. Mayama and K. Tawara . Shionogi Res . Lab.,

Pr idham, T. G. Northern Utilization Res. and Dev. Div., Peor ia ,

Wognicka, Wanda. PaGstwowy Zakaad Higieny. Wars zawa 12, ul.

Shionogi and Co., Ltd., Fukushima-ku, Osaka, Japan.

Illinois, U. S. A.

Chocimska 24, Poland.

Group A- 12

Margalith, P. Labor. of Microbiol. , Dept. of Food and Biotechnology,

Vernon, T . R. Dept. Scientific and Ind. R e s . , P r i v a t e Bag, Auckland

Williams, S . T. Hartley Botan. Labor. , Univ. of Liverpool, Liverpool,

Technion, Haifa, I s rae l .

C1, New Zealand.

England.

Group A- 1 3

k e h a r e k , Z . and Alena kigicovg. Czech. Acad. of Science, Inst. of Microbiol. , Prague 4, Budejovicka No. 1083, Czechoslovakia.

Sanchez-Marroquin, A. Williams, S. T. Hartley Botan. Labor. , Univ. of Liverpool, Liverpool,

Miami 40, Mexico. 18 / D. F., Mexico.

England.

Group B-1

Sanchez-Marroquin, A. Baldacci, E. , G. F a r i n a and R. Locci. 1st. Patol. Vegetale, Univ.

Gauze, G . F . , T . P . Preobrazhenskaya, E.S. Kudrina, T.S. Maximova

Miami 40, Mexico 18 / D . F . , Mexico.

Milano, Milano, Via Celor ia N. 2, Milano, Italy,

and M.A. Sveshnikova. Acad. Med. Sciences of U. S. S. R . , Inst. New Antibiotics, Bolshaia Pirogovskaia 11, Moscow, U. S. S. R.

S Y S T E M A T I C B A C T E R I O L O G Y 28 1

GrouD B- 2

Anderson, Lucia E. and S a r a Wold. Parke , Davis and Co. , Detroit,

C r o s s , T. and A. Maciver. Postgraduate School of Studies in Biological

Johanides, Vera and T e a Blazekovic. Laboratorij za h d u s t r i j s k u mikro-

Michigan, U. S. A.

Sciences, Univ. of Bradford, Bradford, 7 , Yorkshire, England.

biologiju, Tehnolo2ki fakultet, Pierott i jeva 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,

Krasil 'nikov, N. A. Inst. Microbiology. Academy of Sciences, Moscow Nether lands.

B- 133, U. S. S. R.

- Group B-4

Crook, Kenneth E., Jr. and Caro l S. Cassidy. Br i s to l Labs. , Inc.,

Mach, F. Inst. fur Mikrobiologie, Greifswald, Ludwig- Jahn-Str. 15,

Kuznetsov, V. D. USSR Research Inst. for Antibiotics, Moscow, U. S. S. R.

Syracuse, New York, U. S . A.

German Democratic Republic .

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 ,

Szab6, I. Magyar Tudomgnyos Akadgmia, Talajtana ks Agrokgmiai G e r m a n Democratic Republic.

Kutat6 Intgzete, Budapest, Hungary.

Group B-6

kehagek, Z. and Alena &i&vocz.

Mitchell, T . G. T o r r y Research Station, Aberdeen Scotland. Tsyganov, V. A. Res. Inst. of Antibiotics, Leningrad, L-20, U. S . S . R.

Czech. Acad. of Science, Inst. of Microbiol., Prague 4, Budejovicka No. 1083, Cz.echoslovakia.

Group B-8

McClung, Norvel M.* and Gene Michaels. Dept. Microbiol.. Univ. Georgia, Athens, Ga . , U. S.A. *(Present address : Dept. Bot. and Bact., Univ. South Florida, Tampa, Florida. )

Ferm. Div., Nottingham, England:

T . Hasegawa. Takeda Chem. Ind., Ltd., Juso-Higashiyodogawaku, Osaka, Japan.

Spyvee, J. and J. Elliott. Boots P u r e Drug Co., Ltd., Antibiot. and

Nakazawa, K. , M. Shibata, K. Yamamoto, E. Higashide, T. Iwasa and

28 2 I N T E R N A T I O N A L J O U R N A L

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,

Ohara, Y . and H. Nonomura. Faculty of Engineering] Yamanashi Univ., War szawa 12, Poland.

Kofu, Japan.

GrOUD B- 10

Trejo. W. E. R. Squibb and Sons, New Brunswick, New Jersey , U. S. A. B r i n h a n n , Rolf. Organic Chem. Inst., Univ. of Gottingen, Gottingen,

Nishimura, H., M. Mayama and K. Tawara. Shionogi Res. Lab., Germany . Shionogi and Co., Ltd., Fukushima-ku, Osaka, Japan.

Group B-11

Routien, J. B. and Corinne Clevenger. Charles Pfizer and Co. , Inc.

Falczo de Morais, J.0, Inst. de Quimica da Univ. Fed. de Pernambuco,

Okami, Y. Nat'1. Inst. of Health of Japan, Shinagawa, Tokyo, Japan.

Groton, Connecticut, U. S. A.

Recife, Pernambuco, Brazil.

Group B- 1 2

Higgens, Calvin E. and R. E. Kastner. Eli Lilly and Co., Indianapolis,

W i l l i a m s , S. T. Hartley Botan. Labor. , Univ. of Liverpool, Liverpool,

Vernon, T. R. Dept. Scientific and Ind. Res . , Pr iva te Bag, Auckland C1,

Indiana, U. S. A.

England.

New Zealand.

AGKNOW LEDGMENTS

The Subcommittee on Actinomycetes of the Committee on Taxonomy, ASM, participated in the planning of the project, preparation of descriptions and editing of the manuscript. Members of the Subcommittee (other than the authors) during the period covered by this repor t were: T.G. Pridham, E.J. Backus, and W. Grundy.

We a r e grateful t o A.A. Jchida, Jann M. Ichida, Janet Stevens, Mar- gare t Speer, and Louise Hoffhines of the Ohio Wesleyan ISP staff for technical services, for assistance in locating authentic type cultures and descriptive l i terature, and for help in collating data f rom cooperating laboratories.

C. W. Christensen of Difco Laboratories rendered valuable assistance in directing the formulation of dehydrated culture media according to ISP specifications. The technical and financial assistance of Difco Labora- tories, Detroit,. Michigan, in supplying these standardized media to collaborators throughout the world is gratefully acknowledged.


This research was supported by National Science Foundation Grant GB 509 and GB 4365. The Subcommittee on Actinomycetes of the Com- mittee on Taxonomy, ASM, and the Subcommittee on Taxonomy of Actino- mycetes 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 ser ies 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 strain (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 first paper of the ser ies (Gott- lieb and Shirling 1967); descriptive cri teria 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 strains or suggested neotypes a re given in the second paper of the ser ies (Shir- ling and Gottlieb 1968).

This report (Pa r t I11 of the series) includes illustrated descriptions for an additional 100 type strains or suggested neotype strains f rom the genus Streptomyces (including Actinomyces sensu Krasil'nikov) and the genus Streptoverticillium. The second paper of the ser ies (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 three independent collaborators. In r a re instances where significant disagreement occurred in reports from the three laboratories, the culture was either submitted to a fourth collaborator or withdrawn for re- 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. Earlier 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 Pa r t II and Part III of this ser ies a r e deposited a s reference cultures in the American Type Culture Collection, the Centraalbureau voor Schimmelcultures and the USSR Research Insti- tute for Antibiotics. They will also be deposited in an appropriate culture collection in Japan. Cooperative studies characterizing approximately 300 o r mqre additional species a r e in progress o r have been completed. Descriptions for these species will be included in subsequent reports and the type strains will be deposited in the culture collections a s the descriptions a r e published.


ISP CHARACTERIZATIONS OF TYPE STAINS'

Streptomyces aburaviensis Nishmura, Kimura, Tawara, Sa saki, Nakajima, Shimaoka, Okamoto, Shimohira and 1s.ono. Description: N i s h i m u r a e t c. 1957, 205-211. Type strain: S-66 w.). ISP 5033 f r o m Nishimura a s S-66 = ISM 1063.

spore chains moderately long with LO to 50, o r somet imes m o r e than 50, spores 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. 2 ) .

yeas t -mat t agar , oatmeal a g a r , sa l t s - s ta rch a g a r and g lycero l -asparagine a g a r .

Reverse side of colony: Yellowish brown on yeas t -mal 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.

yeas t - i ron 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 , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

Carbon utilization: Growth on carbon utilization media is generally poor. _D-GLucose and _D-fructose and D_-xylose a r e utilized for growth. Reports varied f r o m no growth to t r a c e s of growth on L_-arabinose, sucrose , 1-inositol , D_-mannitol, rhamnose and raffinose.

ISP description by Group B-1. Spore chain morphology: Section Retiflexibiles (Fig. 1 ) . Mature

Color of colony: A e r i a l m a s s color in the Gray co lor -ser ies on

Color in medium: Melanoid pigments a r e not formed in peptone-

S t r eptomy c e s aerocolonigene s Shinobu and Kawato. Description: Shinobu and Kawato 1960a, 212-216. Type strain: 701 w.). ISP 5034 f r o m Shinobu a s 701. ISP description by Group B - 4 .

Morphological charac te r i s t ics : No a e r i a l mycelium i s formed on yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r . In addition to these media, coopera tors found no a e r i a l mycelium o r spore formation on Langeronls weak potato-carrot medium, modified Bennett 's medium, tomato-paste oatmeal agar , soil ex t rac 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 Lose i t s formation ability of a e r i a l mycelia by successive cultures,. . . . I 1 .

Shinobu observed bes 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 charac te r i s t ics : Although ISP collaborators were unable to obtain a e r i a l mycelium on any medium t r ied (see above), Shinobu's original description makes special mention of the formation of secondary colonies on the cottony a e r i a l mycelium on glucose-asparagine a g a r . His i l lustrations suggest that these seconday colonies a r e s imi la r to l lsclerotia ' l repor ted by other au thors (Waksman 1961, 292).

I Type s t r a i n s for LOO Streptomyces spec ies a r e described here . An additional LOO s t r a i n s w e r e descr ibed in an e a r l i e r repor t (og. c i t . ) ; and descriptions for other species a r e in preparation.

Figure 1. 2. aburaviensis. RF spore chains. (a) (X 100) on oatmeal

Figure 2. 2. aburaviensis.

Figure 3 . S. afghaniensis. Spiral spore chains (X 400). Short chains

Figure 4. 5. afghaniensis. Spiny spores; electron micrograph from

agar, 24 days. ’* (b) (X 640) on salts-starch agar, 14 days.

14 day culture on oatmeal agar.

from incomplete spirals or hooks.

14 day culture on oatmeal agar. *

Smooth spores; electron micrograph from

* Sources of illustrations a r e indicated by superscript number following legend for each figure, and a r e listed by number after the bibliography.


Reverse side of colony: No distinctive pigments (pale grayed yellow) on yeas t -mal t agar , oatmeal a g a r , sa l t s - s ta rch a g a r o r g lycero l -aspar - agine a g a r .

yeas t - i ron a g a r , tyrosine a g a r o r peptone-yeast broth. P igments other than t r a c e s of yellow a r e not found in the medium in yeas t -mal t a g a r , oatmeat agar , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

i inositol, -Dmannitol, p f r u c t o s e and rhamnose a r e utilized for growth. sources . No growth o r only t r a c e of growth on raffinose.

Color in medium: Melanoid pigments a r e not formed on peptone-

Carbon utilization: D_-Glucose, L_-arabinose, sucrose , Q-xylose,

Growth on rhamnose is Less abundant than on the other carbon

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.

usually resu l t in hooks, incomplete s p i r a l s o r sp i ra l s with only one o r two turns (Fig. 3) . Short, s t ra ight and flexuous chains of only a few s p o r e s a r e common, but hooks and loops of wide d iameter a s found in typical cu l tures 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 spores may be found on some chains. This morphology i s bes 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 surface: Spiny (Fig. 4).

m e a l a g a r . is in the Yellow color -ser ies . c o l o r - s e r i e s f o r yeas t -mal t a g a r and s a l t s - s t a r c h a g a r . a r e not usually found on glycerol-asparagine a g a r .

to orange o r reddish brown on yeas t -mal t agar , sa l t s - s ta rch 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 formed in peptone-yeast- i ron agar , tyrosine a g a r and peptone-yeast broth. brown pigments a r e found i n the medium in yeas t -mal t agar, 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 . These pigments a r e not pH sensit ive when tested with 0.05 N HCl 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 for growth.

1SP description by Group B-5. Spore chain morphology: Section S p i r a l e s , but shor t spore chains

Color of colony: Aer ia l m a s s color in the Blue co lor -ser ies on oat- Immature aerial mycelium o r mycelium producing few spores

O b s e r v e r s repor ted both Blue and Yellow Mature spores

Reverse side of colony: Grayed yellow on oatmeal a g a r , modified

Orange to r e d o r

Streptomyces atbidofuscue Maeda, Kosaka, Okami and Umezawa 1953. Streptomyces albidofuscus Okami and Umezama Okami, Maeda and Umezawa 1954. Streptomyces pyridomyceticus Okami and Umezawa, 9:~.

The type s t ra in is descr ibed under the name

Actinomyces albocyaneus Krasil 'nikov and Agre. Description: Krasil 'nikov and Agre 1960, 272. Type s t ra in : INMI 679 (2). ISP 5197 f r o m Krasil 'nikov a s INMI 679 = RIA-633. Group B -9.

Spirals, when formed, a r e open and poorly developed. Long chains often show primitive sp i ra l s , t e rmina l sp i ra l s , loops o r hooks charac- te r i s t ic of %cul tures (Fig. 5). Flexuous chains a r e a l s o common.

ISP description by

Spore chain morphology: Section Retinaculiaperti o r Spira les.

Figure 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 agar, 14 days. (b) Open spirals, yeast-malt agar, 14 days. terminal spirals, oatmeal agar, 21 days.

Figure 6. _A. albocyaneus. Spiny spores, short spines; electron micrograph from 10 day culture on yeast-malt agar.

Figure 7. 2. albofaciens. Spiral spore chains (X 1100). agar, 21 days. (b) On glycerol-asparagine agar, 14 days.

Figure 8. 2. albofaciens. Smooth spores; electron micrograph from 14 day culture on glycerol-asparagine agar. '

(c) Flexuous chains and

(a) On oatmeal


Mature spore 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 yeas t -mal 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 shor t and thick (Fig. 6).

s t r a t e mycelium on sa l t s - s ta rch a g a r and glycerol-asparagine a g a r to form I1oidiosporesl1 is repor ted by one o b s e r v e r ; a second observer r e c o r d s ' lchlamydosporesll on oatmeal a g a r and glycerol-asparagine a g a r a f te r 7 days.

Color of colony: Aer ia l m a s s color in 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 sa l t s - s ta rch 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 .

Reverse side of colony: Grayed yellow is modified by green or blue pigments on yeas t -mal t a g a r , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . Only one observer recorded a change in r e v e r s e color f r o m pale green to pale violet by addition of 0.05 N HC1.

Color in medium: Melanoid pigments are not found i n peptone-yeast iron agar , tyrosine a g a r o r tryptone-yeast broth. No pigment i s found in the medium in yeas t -mat t a g a r , s a l t s - s t a r c h a g a r o r g lycero l -asparagine a g a r .

Carbon uti t i za tion : R-G luco s e , &-a rabinos e , sucrose , _D -xy 10s e , - 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 vary f r o m slight growth to good growth for sucrose , &-inositol and raffinose.

Special morphological charac te r i s t ics : Fragmentation of the sub-

Streptomyces albofaciens Thi rumalachar and Bhatt. Description: Thirumalachar and Bhatt 1960, 61-63. Type strain: 27-A w.). ISP 5268 f r o m M. J. Thirumalachar as 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 somet imes appear to a r i s e f r o m a n axial hypha, but t rue whorls typical of verticil late cu l tures a r e not formed. generally LO to 50, o r somet imes m o r e than 5 4 s p o r e s p e r chain. 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. 8).

yeas t -mal t a g a r and glycerol-asparagine a g a r ; White o r Gray 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 .

Reverse side of colony: No distinctive pigments on yeast-malt agar , 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 formed in peptone-yeast- i r o n agar , 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 , oa tmeal agar , sa l t s - s ta rch a g a r o r glycero 1-asparagine a g a r .

Carbon uti Lization: &Glucose, 4 -a rab inose , - inos i to l , _D-mannitol, - I l f r u c t o s e and raffinose a r e utilized for growth. t r a c e of growth on rhamnose. doubtful.

Mature spore chains This

Color of colony: Aer ia l m a s s color in the White c o l o r - s e r i e s on

No growth o r only Utilization of sucrose and _D-xylose is

Streptomyces alboflavus (Waksman and Cur t i s ) Waksman and Henrici . Descriptions: Actinomyces albo-flavus Waksman and Cur t i s 1916, 120 and 128; Actinomyces atbofkivue Waksman 1919, 90-91; Streptomyces alboflavus (Waksman and Cur 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.


Morphological charac te r i s t ics : Typical a e r i a l mycelium i s not formed on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol- asparagine a g a r . Spore chain morphology, spore surface and a e r i a l m a s s color of colony cannot be observed on ISP media. produce sporulating a e r i a l mycelium was noted in an ear ly description of this culture (Waksman 1919, 90). descr ibe 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 charac te r i s t ics : Coremia formation on sa l t s - s ta rch a g a r and glycerol-asparagine agar was recorded by two observers . This s a m e phenomenon was recorded in the original description on Czapekls a g a r (Waksman 1916, 120) : ' I . . . a e r i a l mycelium was found to have a tendency to produce , . . . . . a m a s s of hyphae massed together into a rope, and f r o m this rope fine fi laments coming out in the shape of side branches. and fine rootlets coming out on the side." One ISP observer found straight spore chains within the coremia on glycerol-asparagine agar .

grayed greenish yellow) on yeast-malt agar , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine agar .

Melanoid pigments formed weakly in peptone - yeas t - i ron a g a r and tryptone-yeast broth but not i n tyrosine agar . ments other than melanoids not formed in yeast-malt a g a r , oatmeal a g a r , sa l t s - s ta rch agar o r glycerol-asparagine agar .

- D -mannitol, _D -fructose and raffinose a r e utilized for growth. o r only t r a c e of growth on rhamnose.

Loss of ability to

Ear ly descriptions of the culture

The s t ruc ture looks like the root of a t r e e

Reverse side of colony: No distinctive pigment (grayed yellow o r

Color i n medium: P ig-

Carbon utilization: _D -G luco se , ,L -arabinose, g x y l o se, i - i n 0 sitol, No growth

Utilization of sucrose i s doubtful.

Streptomyces albogriseolus Benedict, Shotwell, Pridham, Linden- f e l s e r and Haynes. Description: Benedict e t a l . 1954, 653-656. Type s t ra in : 7-A = NRRL B -1305 k d . ) . NRRL B-1305. Note: The authenticity of this s t ra in a s the type s t ra in deposited by Dr. R.G. Benedict in the ARS Culture Collection in August, 1951, a s NRRLB-1305 has been verified by C. W. Hesselt ine and T. Pr idham of the ARS Culture Collection, Peor ia , 111. (personal communication). No authentic subcultures f r o m the type s t ra in have hairy spores . At some t ime the culture with hairy spores was apparently mislabelled a s NRRL B-1305 and given limited distribution. spored ISP 5003 (7-A = NRRL B-1305) cor responds to the original description in a l t details . ISP collaborators as ISP 5344. surface, i t differs f r o m the type s t ra in and the original description in carbon utilization. by Benedict e t a t . , u s e s all ISP carbon sources except raffinose and cellulose. carbon sources except _D - glucose and possibly fructose. tion by Group A-2.

flexuous o r spore chains a r e a l s o common. Mature spore chains generally LO to 50 spores p e r chain. mal t agar , oatmeal a g a r and sa l t s - s ta rch agar , but not on glycerol- asparagine agar . Spore surface: Warty. Warts a r e not prominent o r regular and some smooth spores may be fouud (Fig. LO).

ISP 5003 f r o m T. Pr idham a s 7-A =

The warty to smooth

The hairy spored culture was a l so studied by In addition to the difference in spore

The type strain, in agreement with the description

The ha i ry spored s t ra in i s unable to utilize any of the ISP ISP descr ip-

Spore chain morphology: Section Spirales. Spi ra l s a r e open (Fig. $9);

This morphology i s seen on yeast-


Color of colony: Aer ia l m a s s color in the Gray c o l o r - s e r i e s on yeast-matt agar , oatmeal a g a r and sa l t s - s ta rch a g a r .

Reverse side of colony: No distinctive pigments a r e formed (colorless o r yellowish gray on oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine a g a r ; grayed yellowish brown o r olive brown on yeas t - mal t a g a r ) .

yeas t - i ron agar , tyrosine a g a r o r tryptone-yeast broth. No pigment is found in medium in yeast-malt a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

Carbon utilization: _D -Glucose, sucrose , _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 for growth, but growth on sucrose is l e s s abundant than on other carbon sources . No growth or only t r a c e of growth on raffinose.


Streptomyces amakusaens is Nagatsu, Ansai, Ohkuma and Suzuki. Description: Nagatsu et a t . 1963, 207-210. Type s t ra in : 10-101

( i x d .). ISP 5219 f r o m S . Suzuki, Inst. Phys. and Chem. Res . , Tokyo a s LO-101.

chains 10 to 50 o r m o r e s p o r e s p e r chain on yeas t -mal t a g a r and s a l t s - s ta rch a g a r . on oatmea 1 a g a r and glycerol-asparagine agar . Spore surface: Srnooth (Fig. 12).

Color of colony: Aerial m a s s color of mature a e r i a l mycelium i s in the Blue co lor -ser ies on sa l t s - s ta rch a g a r and the Green 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 . a g a r or glycerol-asparagine a g a r .

yellow) on yeas t -mat 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 agar .

yeast-iron a g a r , but not in tyrosine agar . medium in yeast-malt a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r o r glycerol- asparagine a g a r .

This culture does not show good growth with any of the carbon sources tes ted on Pr idham and Gottlieb carbon utilization medium. _D -Glucose is utilized f o r growth. No significant growth occurs on _D -xylose, i -inositol, _D -mannitoL, P f r u c t o s e , rhamnose o r raffinose and only doubtful t r a c e is seen on ,L -arabinose and sucrose .

ISP description by Group B-3. Spore chain morphology: Section Spi ra les (Fig. 11). Mature spore

Sporulating a e r i a l mycelium is poorly developed o r absent

Mature a e r i a l mycelium i s usually not formed on oatmeal

Reverse side of colony: No distinctive pigments (colorless o r pale

Color in medium: Melanoid pigments a r e usually formed in peptone- N o pigments a r e found in the

Carbon uti l iwtion:

Streptomyces ambofaciens Pinnert-Sindico. Description: P inner t -

ISP description by Group

Spore chain morphology: Section Spi ra les (Fig. 13). Open te rmina l

Sindico 1954, 702-707. Type s t ra in : 3486 m.). ISP 5053 f r o m R. Despois, Soci6ZC Rh6ne-Poulenc a s 3486. B -8.

sp i ra l s on long spore chains a l s o s u g g e s t E morphology. spore chains generally LO t o 50 o r m o r e s p o r e s p e r chain. phology i s seen on yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine agar , although sporulating a e r i a l mycelium is not abundant on glycerol-asparagine agar . Spore surface: Smooth to war ty ; surface i r regular i t ies suggesting war ts a r e smal l (Fig. 14).

Mature This m o r -

Figure 9 . 2. albogriseolus. (a, b) Spiral spore chains (X 500) on glycerol-asparagine agar, 7 days.8

F igure 10. S. albogriseolus. Warty and smooth spores ; electron micrograph f r o m 21 day culture on yeas t -mal t agar .

F igure 11. mal t agar, 14 days. '" (b) Tight sp i ra l s , yeast-malt agar , 15 days. ''

Figure 12. S . amakusaensis. Smooth spores ; electron micrograph f r o m 14 day culture on yeas t -mal t agar . l 2

Figure 13. S. ambofaciens. Spi ra l spore chains ( X 800) on sa l t s - s ta rch agar, 1 4 d a y s .

F igure 14. S. ambofaciens. Smooth spores with minor surface i r regu- la r i t i essugges t ing smal l war t s o r spines; electron micrograph f rom 14 day culture on yeast-malt agar . I4

S. amakusaensis. (a) Open and tight sp i ra l s (X 300) on yeast-


Color of colony: Aer ia l m a s s color in the Gray c o l o r - s e r i e s on

Reverse side of colony: No distinctive pigments (colorless to

Dark brown, d a r k blue or a lmost black subs t ra te mycelium pig-

yeas t -mal t a g a r , oatmeal a g a r and sa l t s - s ta rch a g a r .

grayed yellow) on yeast-malt agar , oatmeal a g a r and glycerol-asparagine a g a r . ment i s found on s a l t s - s t a r c h a g a r .

yeas t - i ron agar , tyrosine a g a r or tryptone-yeast broth. may be found in medium in sa l t s - s ta rch a g a r ; no pigment i s in the medium in yeas t -mal t a g a r , oatmeal a g a r or glycerol-asparagine a g a r .

Carbon utilization: _D -Glucose, 4 -arabinose, _D -xylose, i -inositol, - D -mannitol, _D -fructose and rhamnose a r e utilized for growth. growth or only t r a c e of growth on raffinose. doubtful.

This pigment i s not pH sensit ive. Color in medium: Melanoid pigments a r e not formed in peptone-

Yellow pigment

NO Utilization of s u c r o s e i s

Streptomyces aminophilus Wooldridge. Description: The name and oriRinal identification for this species is credited to J. W. F o s t e r by - Oswald e t a t . 1956, 236. This publication contains no description and the original characterizations by F o s t e r have been lost (J. W. F o s t e r , personal communication). The f i r s t available description i s i n Wool- dridge 1957, German Paten t Notation DAS 1000966; s e e a l s o Wooldridge 1960a, Br i t i sh Pa ten t 828, 792 and Wooldridge L960b, U. S . Patent 2,956,925 (Streptomyces coelicotor v a r . aminophilus). NRRL 2390 (Ibld.; L. E. Arnow, W a r n e r - L a d e r t Res. Inst . , personal communication). ISP 5186 f r o m C. W. Hesselt ine a s N R R L 2390. ISP description by Group A-13.

chains 3 t o 10 or somet imes m o r e than 10 s p o r e s per chain. This m o r - phology is seen on yeas t -mal t agar , oatmeal agar , s a l t s - s t a r c h agar and glycerol-asparagine agar , but formation of sporulating a e r i a l mycelium i s poor on oatmeal agar . Spore surface: Smooth (Fig. 16).

yeast-malt a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r ; a e r i a l mycelium poorly developed on oatmeal agar .

yellow) on yeas t -mat t agar , oatmeat agar , sa l t s - s ta rch a g a r or glycerol- asparagine a g a r .

Color i n medium: Melanoid pigments not formed in peptone-yeast- i ron a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigments, o r only t r a c e s of yellow pigment, a r e found in the medium in yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r ,

Carbon utilization: _D -Glucose, _D -xylose, _D -mannitol and _D - f ruc tose a r e utilized for growth. - i -inositol and rhamnose. raffinose.

Type s t ra in :

Spore chain morphology: Section Spi ra les (Fig. 15). Mature spore

Color of colony: Aer ia l m a s s color in the Yellow c o l o r - s e r i e s on

Reverse side of colony: No distinctive pigments (yellow or grayed

No growth o r only t r a c e of growth on Reports vary on utilization of s u c r o s e and

(Waksman and Woodruff) Waksman and Henrici . Description: Actinomyces antibioticus Waksman and Woodruff 1941, 246-249; Streptomyces antibioticus (Waksman and Woodruff) Waksman and Henrici 1948, 942. 1961; R. Gordon, P e r s o n a l Communication). ISP 5234 f r o m S.A. Waksman a s IMRU 3435.

Type s t ra in : IMRU 3435 (Waksman

ISP description by Group B-10.

Figure 15. 2. aminophilus. Spiral spore chains (X 500) on salts-starch

Figure 16. S. aminophilus. Smooth spores; electron micrograph from

Figure 17. S. antibioticus. RF spore chains (X 600). l6

Figure 18. 2. antibioticus. Smootb spores; electron micrograph from 21 day culture on yeast-malt agar. ''

Figure 19. S. antimycoticus. Compact clusters of closed spirals on salts-starch agar, 14 days (X 200)."

Figure 20. S. antimycoticus. Spiny to warty spores in spiral spore chains; electron micrograph from 14 day culture on salts-starch agar. "

agar, 23 days.15

15 day culture on yeast-malt agar.

-


Spore chain morphology: Section Rectiflexibiles (Fig. 17). Mature spore 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. agar , oatmeal agar , s a l t s - s t a r c h a g a r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 18).

yeast-malt agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r ; mature a e r i a l mycelium poorly developed on oa tmeal agar .

Reverse side of colony: No distinctive pigments (grayed yellow- brown on yeas t -mal t agar , grayed yellow o r d a r k greyish greenish yellow on oatmeal agar , s a l t s - s t a r c h a g a r , glycerol-asparagine a g a r ) ; subs t ra te mycelium pigment is not a pH indicator.

Color in medium: Melanoid pigments a r e produced in peptone-yeast- i ron a g a r , tyrosine a g a r and peptone-yeast broth. o r only t r a c e of yellow, in yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

- D-fructose and rhamnose a r e utilized for growth. t r a c e of growth on sucrose and raffinose. doubtful.

This morphology is observed on yeas t -mal t

Color of colony: Aer ia l m a s s color in the Gray co lor -ser ies on

No pigment in medium,

Carbon utilization: ,D -Glucose, &-arabinose, i - inos i to l , _D-mannitol, No growth o r only

Utilization of _D-xylose i s

Streptomyces antimycoticus Waksman, Descriptions: Streptomyce s 9 Leben, S tesse l and Keitt 1952, 159-168; Streptomyces antimycoticus Waksman 1957, 799. Type s t ra in : A158 m). ISP 5284 f r o m C. W. Hesselt ine a s NRRL 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 and glycerol-asparagine a g a r (Fig. 19); mature spore chains of generally LO to 50 spores p e r chain on these media. warty. short , thick spines and war ts (Fig. 20).

yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

grayed greenish yellow) on yeas t -mal t agar , oatmeal a g a r , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

Color i n medium: Melanoid pigments a r e not produced on peptone- yeast-iron agar , tyrosine a g a r o r tryptone-yeast broth. No pigment i s found in medium i n yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

- i -inositol, _D-mannitol, _D-fructose, rhamnose and raffinose a r e utilized for growth. abundant than OD the other carbon sources .

Spore surface: Spiny to Surface i r regular i t ies on s p o r e s a r e intermediate between very

Color of colony: Aer ia 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

Reverse side of colony: No distinct pigment (pale grayed yellow o r

Carbon utilization: _D-Glucose, &-arabinose, sucrose , _D-xylose,

Growth on _L-arabinose and 2-xylose is generally less

Streptomyces a rab icus Shibita, Nakazawa, Miyake, Inoue and Okabori. Description: Shibita et a l . 1957, 32-37. Type s t ra in : 6762 m.). description by Group B-8.

sa l t s - s ta rch a g a r (Fig. 2La). and glycerol-asparagine a g a r may appear to be RF morphology o r may

ISP 5252 f r o m K. Nakazawa a s IF0 3406 = 6762. ISP

Spore chain morphology: Section Spi ra les on oatmeal a g a r and Short spore chains on yeast-malt a g a r


show only incomplete sp i ra l s or hooks (Fig. 21b). on oatmeal a g a r and sa l t s - s ta rch a g a r generally LO to 50 or m o r e spores p e r chain; shor te r spore chains are 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 shor t and blunt (Fig. 22a); smooth s p o r e s may a l s o be found (Fig. 22b).

yeas t -mal t agar , oatmeal a g a r and sa l t s - s ta rch a g a r ; Red co lor -ser ies on glycerol-asparagine a g a r . ing neutral g r a y s to light grayish reddish brown (5fe) for the Gray color- s e r i e s and grayish yellowish pink (5dc) f r o m the Red co lor -ser ies for glycerol-asparagine a g a r .

Reverse side of colony: Grayish yellowish brown on yeast-malt a g a r ; co lor less to pale yellowish gray o r brown on aa tmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . a pH indicator.

i ron agar, ty ros ine agar , o r tryptone-yeast broth. T r a c e of yellow pigment may be found in the medium i n oa tmeal agar or sa l t s - s ta rch agar or m a y be absent.

- D-mannitol, _D-fructose and rhamnose a r e utilized for growth. growth or only trace of growth on raffinose. doubtful.

Mature spore chains

Color of colony: A e r i a l m a s s color i n the Gray co lor -ser ies on

Observers selected color tabs represent -

Substrate mycelium pigment is not

Color in medium: Melanoid pigment is not produced in peptone-yeast-

This pigment is not pH sensitive. Carbon utilization: D- Glucose, 2 -arab inose , D-xylose, i=inositol,

Utilization of s u c r o s e is No

Streptomyces argenteolus Fr ied , Per lman, Langlykke and Titus. Descriptions: Actinomycete spec ies ATCC 11,009 P e r l m a n e t at. US. Patent 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 as MD 2428 = ATCC 11,009.

Spore chain morphology: Section Spirales (Fig. 23). This culture is described as Itnot forming loops o r spirals" i n the original descr ip- tions appearing in the patents cited above. However, the type s t ra in was observed to produce s p i r a l s in subsequent studies by Pr idham e t at., 1958 and by T r e s n e r et at., 1961. Mature spore chains generally 10 t o 50 o r m o r e spores p e r chain on yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 24).

yeast-malt agar , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine agar .

Reverse side of colony: Colorless to grayish yellowish green o r gray on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine a g a r ; subs t ra te mycelium pigment is not a pH indicator.

Color in medium: Melanoid pigments not formed in peptone-yeast- i r o n agar , tyrosine a g a r or tryptone-yeast broth. No pigment found in medium i n yeast-malt agar , oatmeal agar , sb l t s - s ta rch a g a r or g Lycerol-asparagine agar .

- D-fructose and rhamnose are utilized for growth. t r a c e of growth on sucrose, i - i n o s i t o l and raffinose.


--

Color of colony: Aer ia l m a s s color i n the G r a y co lor -ser ies on

Carbon utilization: _D-Glucose, &-arabinose, _D-xylose. _D-mannitol, N o growth or only

Figure 21. S. a rab icus . (a) Spi ra l spore chains ( X 300) on s a l t s - s t a r c h agar , (b) Short, flexuous chains with some hooks and incomplete sp i ra l s ( X 800) on yeast-malt agar , 14 days. l 3

a r e thick a n d o r rounded.19 (b) Smooth spores of S. a r a b i c u s ; electron micrograph f r o m 1 4 day culture on yeas t -mal t agar . I4

F igure 22 . S. arabicus. (a) Spiny t o warty spores ; sur face i r regular i t ies

- Figure 23 . 2. a r enteolus. Spi ra l spore chains ( X 600) on yeas t -mal t

F igure 24. S. argenteolus. Smooth spores ; electron micrograph f r o m

agar , 14 days. --%---

14 day culture on oatmeal a g a r . lo


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) Pr idham e t at. 1958, 67. ISP 5138 f r o m T.P. Preobrazhenskaya a s 10369/58.

Spore chain morphology: Section Retinaculiaperti o r Spirales. Spi ra l s a r e open and often a r e i r r e g u l a r and poorly developed 25a, b). Long spore chains of the = type and flexuous chains a r e common (Fig. 25c). a r e moderately long with m o r e than.10 spores p e r chain on oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 26).

co lor -ser ies on yeast-malt agar , sa l t s - s ta rch a g a r and glycerol- asparagine a g a r ; both co lors may appear on the s a m e medium. observer noted an increase in gray a e r i a l color between 14 and 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. frequently chosen were 2ec (yellowish gray) f r o m the Gray co lor -ser ies and 5ge (tight grayish reddish brown) f r o m the Red co lor -ser ies .

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 sa l t s - s ta rch agar). Substrate mycelium pigment is not a pH indicator.

Color in medium: Melanoid pigments a r e produced in peptone-yeast- i 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 in yeast-malt agar , oatmeal agar , sa l t s - s ta rch 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.

ISP description by Group B - 6 .

(Fig.

Mature spore chains may be slow to develop; they

Color of colony: Aer ia l m a s s color i s in both the Red and the Gray

One

The color tabs most

Reverse side of colony: No distinctive pigment (grayed yellow on

St r ep t omyce s a u r e ofa ciens Dugga r . De s c ription: Dug ga r 1948, 17 7 - ISP 5127 f r o m E. Backus, Lederle Labs., Amer .

ISP description by Group A-12. Spore chain morphology: Section Retinaculiaperti but chains r e p r e -

This morphology is seen

181; U. S. Patent 2,482,055 Sept. 1949. Type strain: A-377 = NRRL 2209 k d . ) . Cyanamid Co. a s A-377 = NRRL 2209.

sentative of Section K a r e a l s o common (Fig. 27). generally LO to 50 o r m o r e spores p e r chain. on yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine agar . Spore surface: Smooth (Fig. 28).

mat t agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar .

orange-yellow o r brown on yeast-malt agar , and grayed yellow o r green- i s h yellow on oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar ) .

i ron agar , tyrosine a g a r o r tryptone-yeast broth. in medium in yeast-malt a g a r , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine agar .

- D-fructose a r e utilized f o r growth. on&-inositol, _D-mannitol, rhamnose and raffinose.

Mature spore chains

Color of colony: A e r i a l mass color i n the Gray co lor -ser ies on yeast-

Reverse side of co tony: No distinctive pigments (grayed yellow,

Substrate myce Lium pigment i s not a pH indicator. Color i n medium: Melanoid pigments not formed in peptone-yeast-

No pigment is found

Carbon utilization: D_-Glucose, &-arabinose, sucrose, _D-xylose and No growth o r only t r a c e of growth

Figure 25. - A. aurant iogr i seus . (.a) Incomplete s p i r a l s o r hooks ( X 500) on yeas t -mal t a g a r , 23 days. l5 (b) Spi ra l s (X 700) on yeas t -mal t agar , 2 1 days." (c) Flexuous spore chains ( X 350) on yeas t -mal t agar, 18 days."

F igure 26. A. aurant iogr i seus . Smooth s p o r e s ; e lec t ron micrograph f r o m 1 8 2 a y cu l ture on yeas t -mal t a g a r . "

Figure 27. S. aureofaciens. RA spore chains ( X 240) on oa tmeal agar , 14 days?

F igure 28. S. aureofaciens. Smooth s p o r e s ; electron micrograph from 14 day culture on oatmeal agar. ''

Figure 29. 2. baarnensis. Smooth s p o r e s ; electron micrograph from 14 day culture on glycerol-asparagine a g a r . '

-

S Y S T E M A T I C B A C T E R I O L O G Y 299

(Duchg) Pridham, Hesselt ine and Benedict. Descriptions: Actinomyces v i r id i s Duchd 1934, 311-317; Actinomyces baarnens is Duchd 1934 = Actinomyces v i r id i s Gougerot, B lum and Duchk 1934, 376. Type strain: Dreyfus 472 (G.A. deVries, CBS, .-

personal communication). ISP 5232 f r o m G.A. deVr ies as Dreyfus 472. ISP description by Group B-1.

Spore chain morphology: This s t ra in has apparently lost the ability to produce good sporulating a e r i a l mycelium. Spore chains, when found, were straight (Section Rectiflexibiles) and usually contained m o r e than LO spores p e r chain. Spore surface: Smooth (Fig. 29).

sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r is inadequate for de te r - mination of a e r i a l m a s s color. The original description of Duchd h.2.) descr ibes e a r l y appearance of abundant greenish aer ia l myce lium on comparable media.

Reverse side of colony: No distinctive pigment; yellow-brown on yeast-malt agar , light grayish yellow on oatmeal agar , sa l t s - s ta rch agdr and glycerol-asparagine agar .

i ron agar , tyrosine a g a r o r tryptone-yeast broth. No pigment is found i n the medium i n yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r o r g lycero 1 -asparagine agar.

- D-fructose and rhamnose a r e utilized for growth. t r a c e of growth on raffinose. doubtful.

Color of colony: Aer ia l mycelium on yeast-matt a g a r , oatmeal agar ,

Color i n medium: Melanoid pigment is not formed in peptone-yeast-

Carbon utilization: 2-Glucose, &-arabinose, _D-xylose, g-mannitol , No growth o r only

Utilization of sucrose andi - inos i to l is

Streptomyces bellus Margalith and Bere t ta . Description: Margalith and Bere t ta 1960, 189-195. ISP 5185 f r o m L.G. Silvestri , Lepetit S.p.A. as Lep. M. A/870. ISP description by Group A-13.

Spore chain morphology: Section Spirates. Short spore chains may f o r m incomplete spirals, hooks, and loops of smal l d iameter o r flexuous chains (Fig. 30). Hooks and loops a r e of smal l d iameter 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, spores p e r chain on yeast-malt agar , oatmeal a e a r , sa l t s - s t a r c h a g a r and glycerol-asparagine agar . Spore surface: Spiny (Fig. 31).

Color of colony: Aer ia l m a s s color in the Blue co lor -ser ies on oatmeal a g a r and sa l t s - s ta rch a g a r ; White o r Blue co lor -ser ies on yeast- mat t a g a r and glycerol-asparagine agar .

Reverse side of colony: Substrate color is modified by red (orange) pigment on yeast-malt a g a r and glycerol-asparagine agar and by red o r blue pigments on oatmeal a g a r and sa l t s - s ta rch a g a r . These pigments a r e not pH indicators.

Color in medium: Melanoid pigments a r e formed in peptone-yeast- i r o n agar , tyrosine a g a r and tryptone-yeast broth. may be found i n the medium in glycerol-asparagine a g a r ; it is not a pH indicator. oatmeal a g a r o r sa l t s - s ta rch agar .

Carbon uti t i za tion: _D -G lucos e, k-ara binos e, sucro se, E-xy Lose - i-inositot, 2-mannitol , g.fructose, rhamnose and raffinose are utilized for growth.

Type strain: A/870 ( x d . ) .

Spi ra l s a r e best developed on sa l t s - s ta rch agar .

-

T r a c e of r e d pigment

Pigments a r e not found in the medium i n yeast-malt agar ,


Streptomyces bikiniensis Johnstone and Waksman. Descriptions: Johnstone and Waksman 1947, 294; Johnstone and Waksman 1948, 317- 326. Type strain: IMRU 3514 (S.A. Waksman and R. Gordon, personal communication). ISP 5235 f r o m S.A. Waksman a s IMRU 3514. ISP description by Croup B-9.

spore 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 is s e e n on yeas t -mal t agar , oa tmeal agar , sa l t s - s ta rch 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 yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and g lycero l -aspar - agine a g a r .

grayed yellowish brown) on yeas t -mat t a g a r , oa tmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

i ron a g a r , tyrosine a g a r o r tryptone-yeast broth. ment, o r no pigment may be found in medium i n tryptone-yeast broth, yeas t -mal t a g a r , oa tmeal a g a r , sa l t s - s ta rch a g a r and g lycero l -aspar - agine a g a r ; this pigment is not a pH indicator.

a r e utilized f o r growth. Utilization of _L-arabinose, sucrose , i - inos i to l and raffinose if doubtful.

Spore chain morphology: Section Rectiflexibiles (Fig. 32). Mature

Reverse side of colony: No distinctive pigment (grayed yellow to

Color in medium: Melanoid pigment is not produced in peptone-yeast- T r a c e of yellow pig-

Carbon utilization: _D-GLucose, _D-xylose, D_-mannitol and _D-fructose N o growth o r only t r a c e of growth on rhamnose.

Actinomyces biverticil tatus Preobrazhenskaya. Description: Preobrazhenskaya ia Gauze e t a t . 1957, 75-76. Type s t ra in : 10204/54 (G. F. Gauze, personal communication; C. F. Gauze cited by Cottlieb, 1968, 20). biverticil latus (Preobrazhenskaya) Pridham gal,, 1958, 72; Strepto - ver t ic i l l ium biverticil latus (sic) (Preobrazhenskaya) Baldacci 1958,25; Streptoverticil l ium biverticil latum (Preobrazhenskaya) B a ldacci, Farina and Locci 1966, 157. be identical with Streptomyces h i rosh imens is Shinobu 1955, personal communication, 1964. ISP description by Group B -3.

ver t ic i l la te (biverticil late) spore chains (Fig. 34). generally contain 10 o r m o r e s p o r e s p e r chain. on yeas t -mal t agar , oatmeal a g a r , s a l t s - s t a r c h a g a r and g lycero l -aspar - agine a g a r . Spore surface: Smooth (Fig. 35).

mal t agar , oatmeal agar , s a l t s - s t a r c h a g a r and glycerol-asparagine agar . s e r i e s on yeas t -mal t a g a r and sa l t s - s ta rch agar).

Reverse side of colony: Reverse color is r e d on yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r ; this pigment is pH sensitive, changing f r o m r e d t o orange- red with addition of 0.05 N NaOH and f r o m r e d t o violet-red o r blue with addition of 0.05 N HC1.

Color in medium: Melanoid pigments are formed i n l e s s than 2 days in peptone-yeast-iron a g a r and tryptone-yeast broth, but m o r e slowly in tyrosine agar . P igments other than melanoids a r e not formed i n yeas t - mal t a g a r , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar .

ISP 5272 f r o m G. F. Gauze as 10204/54. Streptomyces

The original au thors now consider this to

Spore chain morphology: Section Verticil lati with umbellate-mono- Mature spore chains

This morphology i s seen

Color of colony: Aer ia l m a s s color in the Red c o l o r - s e r i e s on yeas t -

(One observer , only, placed this culture in the Violet co lor -

F i g u r e 3 0 . S. bellus. (a) Spiral spore chains on s a l t s - s t a r c h agar , 21 days. 23-(b)mplete sp i ra l s , hooks and loops (X 550) on s a l t s - s ta rch agar , 24 days. l 5

micrograph f r o m 24 day culture on yeas t -mal t a g a r . 24

14 days: 25

14 day cu l ture on yeast -malt a g a r . ' morphoiogy (X 6 0 0 ) . (a) O n y e a s t - m a l t agar , 7 days; (b) On glycerol- asparagine agar , 14 days. 2o

14 day culture on s a l t s - s t a r c h a g a r . l o

Figure 31. S. bellus. Spiny spores , chromium shadowed; electron

Figure 3 2 . S. bikiniensis. R F spore chains (X 240) on yeast-malt agar ,

F igure 3 3 . S. bikiniensis. Smooth spores ; electron micrograph f r o m

Figure 34. A. biverticil latus. BIV (umbellate monoverticillate)

- -

Figure 35. 4. biverticil latus. Smooth spores ; Blectron micrograph f r o m


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 oni - inos i to l and 2- f ruc tose is much less abundant than on _D- glucose. No growth o r only t r a c e of growth on - L-arabinose, sucrose , _D-xylose, _D-mannitol, r hamnose and raffinose.

Streptomyces bobili (Waksman and Cur t i s ) Waksman and Henrici . Descriptions: Actinomyces bobili Waksman and Cur t i s 1916, 121 and 127; Waksman 1919, 100-102; Streptomyces bobiliae (&)(Waksman and Cur t i s ) Waksman and Henrici 1948, 937. Type s t ra in : IMRU 3310 (Waksman 1961, 182). ISP 5056 f r o m S.A. Waksman as IURU 3310. ISP descr ip- tion by Group B -11.

mycelium is not produced a t 27'c o r 37'C on yeas t -mal 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 observer . found some spots of sporulating a e r i a l mycelium on sa l t s - s ta rch a g a r . One collaborator (J.B. Routien) found sp i ra l spore 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 basa l s a l t s medium for carbon utilization enriched with xylose, raffinose o r glucose (Fig. 36). Incubation was a t 37°C. Spi ra l s were most abundant when xylose was the carbon source . notes the absence of t rue a e r i a l mycelium o r s p o r e s on Czapek's medium. The description published in 1919 (Waksman, 9. e.) notes sp i ra 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 ta rch a g a r plate and potato plug. Waksman. descriptions. Spore sur face : Smooth (Fig. 37).

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 , Sparse a e r i a l mycelium, when produced, is white.

Reverse side of colony: Substrate mycelium color may be grayish yellow, o r may be modified with r e d to reddish gray o r pink, reddish brown, o r reddish orange. The subs t ra te 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 . yeas t - i ron agar , tyrosine a g a r o r tryptone-yeast broth. a r e found in medium i n yeast-matt agar , oa tmeal a g a r , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

Carbon utilization: _D-Glucose, _L-arabinose, sucrose , _D-xytose, - i -inositol, 2 - f ruc tose , rhamnose and raffinose are utilized for growth. No growth o r only t r a c e of growth on 2-mannitol .

Spore chain morphology: Section Spirales. Sporulating a e r i a l

Two o b s e r v e r s

The f i r s t description by Waksman (1916 9. cit .)

A e r i a l mycelium was not produced on 7 other solid media used by The present culture s e e m s to conform well to the ear ly

Color of colony: Aer ia l m a s s color cannot be observed on yeas t -mal t

Color in medium: Melanoid pigments a r e not produced in peptone- N o pigments

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 ra in : T-3018 w.). ISP 5010 f r o m H. T r e s n e r as T-3018. Group B-LO.

(Fig. 38). Spore chains a r e shor t and poorly developed or absent on yeas t -mal t a g a r , oatmeal a g a r and glycerol-asparagine agar . 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 is poorly developed on most media with

ISP description by

Spore chain morphology: Section Spi ra les on sa l t s - s ta rch a g a r .

The

F i g u r e 3 6 . S. bobili. Sp i ra l spore chains (X 50) on Pr idham and Gottlieb

F igure 37. S. bobili. Smooth spores ; electron micrograph ( X 2, 700) carbonGti1ization medium plus raffinose. 36

f r o m 13 day culture on Pr idham and Gottlieb carbon utilization medium plus raffinose. 36

-__ .

Figure 3 8 . S. calvus. Spi ra l spore chains on s a l t s - s t a r c h agar . l 6

Figure 39. S . calvus. Spiny to hairy spores ; electron micrograph f r o m 21 day F u l h G n sa l t s - s ta rch agar .”

F igure 40. A. candidus. R F spore chains ( X 8 7 5 ) . (a) Flexuous a e r i a l chains on yeas t -mal t aga’r, 22 days; (b) Long chains on glycerol- asparagine agar surface, 22 days. 27

14 day culture on oatmeal a g a r . l 2

- -

- - -

Figure 41. A . candidus. Smooth s p o r e s ; electron micrograph f r o m - -


best sporutation and sp i ra l development on s ta rch containing inorganic sa l t s media. Spore surface: Spiny to hairy (Fig. 39).

Color of colony: Aer ia l mycelium is inadequate for color de te rmina- tion on most media. on yeas t -mat t a g a r o r s a l t s - s t a r c h agar , i t is in the Gray co lor -ser ies .

g ray ish yettowish brown) on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

Color in medium: Melanoid pigments a r e not found in peptone-yeast- i ron a g a r , tyrosine a g a r o r tryptone-yeast broth; no pigment is found in medium in yeas t -mal t a g a r , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol- asparagine a g a r .

i - -inositol, _D-mannitol, 2 - f ruc tose , rhamnose and raffinose a r e a l l utilized for growth.

When mature sporulating a e r i a l mycelium i s formed

Reverse side of colony: No distinctive pigments (colorless o r

Carbon utilization: _D-GLucose, _L-arabinose, sucrose , _D-xylose,

Actinomyce s candidus Krasil 'nikov. Descriptions : K r a s i llnikov 1941, 49; Krasil 'nikov 1949, LOO. Type s t ra in : 5855/54 (neotype, P r e o b r a zhenskaya, persona 1 communication ; Kra sill nikov, personal communication). 94. ISP 5141 f r o m T . P . Preobrazhenakaya a s 5855/54. ISP descr ip- tion by Group A-3.

Spore chain morphology: Section Rectiflexibiles. Mature spore 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 sa l t s - s ta rch a g a r and glycerol-asparagine agar . and oatmeat a g a r . Spore surface: Smooth (Fig. 41).

yeas t -mal t a g a r , oatmeat a g a r , sa l t s - s ta rch a g a r and g lycero l -aspar - agine a g a r , although a e r i a l mycelium may be poorly developed on yeas t - m a Lt a g a r and oatmea 1 a g a r .

Reverse side of colony: No distinctive pigments (co lor less or v e r y pate grayed yellow) on yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

i ron agar , tyrosine a g a r o r tryptone-yeaat broth. P igments a r e not formed in medium in yeas t -mal t agar , oatmeal a g a r , sa l t s - s ta rch 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 for growth. growth on sucrose , i - inos i to l and raffinose.

Streptomyces candidus (Krasil 'nikov) Waksman 195 3,

Sporulation may be poor on yeas t -mal t a g a r

Color of colony: Aer ia l m a s s color in the White c o l o r - s e r i e s on

Color in medium: Melanoid pigments not formed in peptone-yeast-

No growth o r only t r a c e of

Streptomyces capreolus Higgens. Description: Higgens & S t a r k -- e t at . 1962, 596-606. Lively, Lilly Res. Labs. , personal communication). ISP 5225 f r o m

Type s t ra in : M48-E2655 = NRRL 2773 (D.H.

D.H. Lively as M48-EZ655 = NRRL 2773.

of 50 o r m o r e spores p e r chain on mature cu l tures (Fig. 42) on yeas t - 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 . Spore surface: Smooth (Fig. 43).

c o l o r - s e r i e s a f t e r 14 to 21 days. White co lor -ser ies .

ISP description by Group B-2. Spore chain morphology: Section Rectiflexibiles with long spore chains

Color of colony: Aer ia l m a s s color i s usually in the Red o r Yellow Immature a e r i a l mycelium is in the

Color tabs selected by o b s e r v e r s for 21 day cu l tures


included 2ba (pale yellow) f r o m the Yellow color -ser ies and 3ca (pale orange yellow) to 5cb (grayish yellowish pink) f r o m the Red color- s e r i e s . These colors a r e observed on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

on yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine a g a r ; this pigment is not a pH indicator.

yeast-iron agar , tyrosine a g a r o r tryptone-yeast broth. Yellow or yellow orange pigment i s found in the medium in yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and usually glycerol-asparagine a g a r ; this pigment is not a pH indicator.

for growth. Reports vary on utilization of _D-xylose, _D-mannitol and - D-fructose, although in each case, two o r th ree 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 sugars a r e not utilized.

Reverse side of colony: Grayed yellow to orange yellow o r orange


Carbon utilization: _D-GLucose, _L-arabinose and i - inos i to l a r e 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 . ) = Pf izer 6563 (J. Routien, Chas. Pf izer and Co., personal communication). 5258 f r o m J. Routien a s 6563 = ATCC 12,476. B-5.

chains of 3 to LO spores occur in dense c lus te rs s o that morphology i s difficult to determine (Fig. 44a). incomplete sp i ra l s (Fig. 44b). c lus te rs but mos t chains a r e too short to f o r m t rue sp i ra l s (Fig. 44c). Short chains a r e a l s o not typical of seen on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine agar . Davisson and Finlay (%.At.). Spore surface: Smooth (Fig. 45).

yeast-malt agar , oatmeal a g a r and glycerol-asparagine a g a r ; Gray o r Green co lor -ser ies on sa l t s - s ta rch a g a r . medium gray to olive gray o r grayish olive (see tabs 2ih o r 2ge in Gray co lor -ser ies and 1 1/2ge in Green co lor -ser ies ) .

Reverse side of colony: Colorless o r grayish yellow to light grayish olive o r olive brown on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . Substrate mycelium pigment is not pH sensit ive.

Color in medium: Melanoid pigments a r e not found in peptone-yeast- i ron agar , tyrosine a g a r o r tryptone-yeast broth; no pigment o r only t r a c e of yellow pigment in medium in yeast-malt a g a r , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar .

Carbon ut lization: _D -Glucose, _D -mannito 1, _D -fructose and raffino s e a r e utilized for growth. No growth or only t r a c e of growth on _L-arabinose, sucrose , _D-xylose, i- inositol and rhamnose.

ISP ISP description by Group

Spore chain morphology: Section Rectiflexibiles. Very shor t spore

Short chains often f o r m hooks o r Tight sp i ra l s may occur in the dense

cultures. This morphology is

It is representative of morphology described by

Color of colony: Aer ia l m a s s color in the Green co lor -ser ies on

The color on a l l media i s

Streptomyce s ce 1 l o staticu s Hamada . Description: Hamada 1958, 173-179. Univ. a s E-150.

Type strain: E-150 w.). ISP 5189 f r o m N. Ishida, Tohoku ISP description by Group A-13.

Figure 42. S. capreolus. Long RF spore chains ( X 380) on Waksman

Figure 4 3 . S. capreolus. Smooth spores ; electron micrograph from

Figure 44. S. catenulae. (a ) Large, dense c lus te rs of spore chains

s t a r c h :gar B, 21 days."

2 1 day culture on s a l t s - s t a r c h a g a r . 2 9

(X 400);n m r c h agar , 14 days. chains, some of which appear to be bent into hooks o r par t ia l s p i r a l s ( X 400) on s a l t s - s t a r c h agar , 14 days. (c ) Short, twisted spore chains (X 300) on s a l t s - s t a r c h agar , 14 d a y s . 4

F i g u r e 45. S. catenulae. Smooth s p o r e s ; electron micrograph f r o m 32 day culture on yeas t -mal t a g a r . 30

(b) Clus te rs of shor t spore

-~


Spore chain morphology: Section Spirales (Fig. 46). Mature spore chains LO to 50 spores p e r chain; longer chains a r e often observed. This morphology is seen on yeast-malt agar , oatmeal agar , sa l t s - s t a r c h agar and glycerol-asparagine agar . Spore surface: Spiny (Fig. 47).

s e r i e s on yeast-malt a g a r , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine agar . grayish yellowish brown) and 5ge o r 5dc (grayish yellowish pink) color tabs of Tresner -Backus color wheels.

yellow o r light yellow brown) on yeast-malt agar , oatmeal agar , sa l t s - s t a r c h agar o r glycerol-asparagine agar .

i ron agar , 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 agar , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine agar .

Carbon utilization: 2-Glucose, &-arabinose, sucrose, _D-xylose, - i -inositol, D_-mannitol, 2 - f ruc tose , rhamnose and raffinose a r e utilized for growth.

Color of colony: Aer ia l m a s s color in the Red (or Gray) color-

Charac te r i s t ic color is usually between 3ge (light

Reverse side of colony: No distinctive pigment (charac te r i s t ic grayed

Color in medium: Melanoid pigments a r e formed in peptone-yeast-

Streptomyces chibaensis Suzuki, Nakamura, Okuma and Tomiyama. Description: Suzuki et a t . 1958, 81-83. Type strain: 77-SN-2 (Suzuki, personal communication). ISP 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). spore chains a r e straight and many a r e flexuous o r curved to f o r m hooks

ISP 5220 f r o m S. Suzuki a s 77-SN-2.

Some

o r par t ia l sp i ra l s (Fig: 48b). o r 20 spores p e r chain) for this culture to be placed in section 48b). sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . Smooth (Fig. 49).

co lor -ser ies on yeast-malt a g a r , oatmeal a g a r and sa l t s - s ta rch a g a r ; Gray or Yellow color -ser ies on glycerol-asparagine agar . Color tabs selected f r o m the Gray co lor -ser ies were yellowish gray o r grayish yellowish brown.

Reverse side of colony: No distinctive pigments (grayed greenish yellow, yellow o r r a r e l y orange-yettow) on yeast-matt agar , oatmeat agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r ; substrate mycelium pigment is not a pH indicator.

i ron 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, sucrose , D_-xylose, - i -inositot, 2-mannitol , _D-fructose, rhamnose and raffinosc a r e a l l utilized for growth.

Moat spore chains a r e too shor t (3 to LO

(Fig. This morphology is seen on yeast-malt agar , oatmeal agar ,

Spore surface:

Color of colony: Mature a e r i a l m a s s color is usually in the Gray

Color in medium: Melanoid pigments a r e not found in peptone-yeast-

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.

Figure 46. S . cellostaticus. Spi ra l spore chains forming tight coils. - (a ) On yeas t -mal t agar , 21 days.23 (b) On s a l t s - s t a r c h agar , 2 2 days ( X 500). l 5

13 day culture on Gauze ' s medium No. 1.

(a) Spiral chains; (b) Flexuous chains."

2 1 day cu l ture on oatmeal a g a r . "

F i g u r e 47. S. cellostaticus. Spiny s p o r e s ; electron micrograph f r o m

Figure 48. S. chibaensis. Spore chains (X 800) on oa tmeal a g a r , 14 days:

F igure 49. S. chibaensis. Smooth spores ; electron micrograph from


Gauze cited by Gottlieb 1968, 20). (Preobrazhenskaya, Ryabova and B linov) Pr idham e t at . 1958, 68. ISP 5273 f r o m G. F. Gauze a s 13638/58. ISP Description by Group B-4.

Streptomyces chromofuscus

Spore chain morphology: Section Spi ra les (Fig. 50). Mature spore chains moderately Long with LO to 50, o r often m o r e than 50, spores p e r chain. This morphology is 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 agar . Spiny (Fig. 51).

yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

Reverse side of colony: No distinctive pigment (grayed yellow to olive brown) on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and g lyce r o 1 -a sparagine a g a r .

Color i n medium: Melanoid pigments a r e formed in peptone-yeast- i ron a g a r and tryptone-yeast broth, but not i n tyrosine a g a r . other than melanoids o r faint t r a c e s of yellow are not found in the medium in yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

- D-mannitol, _D-fructose and rhamnose a r e utilized for growth. v a r y on utilization of raffinose andi - inos i to l . of growth on sucrose.

Spore surface:

Color of colony: Aer ia l m a s s color in the Gray co lor -ser ies on

Pigments

Carbon uti lization: 2 -G lucose, _L-arabino s e, ,D --lo se, i - i n o sitol,

NO growth o r only t r a c e Reports

Streptoverticil l ium cinnamoneum (Benedict, Dvonch, Shotwe 11, P r i d - h a m and Lindenfelser) Baldacci, Far ina , 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); Streptoverticil t ium cinnamoneus (81c) (Benedict e t at. ) Baldacci 1958, 25; ,SJreptoverticillium cinnamoneum (Benedict et at. ) -- emend. hobis Baldacci, Far ina , Locci 1966, 158. Type strain: NRRL B-1285 (T. Pridham, personal communication). P r i d h a m a s NRRL B-1285.

section Verticil lati , biverticil late) (Fig. 52). 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 is seen on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 53).

yellowish pink) on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar .

Reverse side of colony: No distinctive pigment (grayish yellow to yellow-brown o r brown) on yeast-malt agar , oatmeal agar , sa l t s - s t a r c h a g a r and glycerol-asparagine agar .

i ron agar , tyrosine a g a r o r tryptone-yeast broth. No pigment i s found in the medium i n yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r o r g Lycerol-asparagine agar .

Carbon utilization: Vegetative growth is generally poor on Pridham and Gottlieb's carbon utilization medium plus _D- glucose. collaborators var ied f r o m no growth to slight growth with other carbon sources , but good growth was not observed on any of the following: - L-arabinose, sucrose , _D-xylose, g-inositol, _D-mannitol, 2 - f ruc tose ,

ISP 5005 f r o m T. ISP description by Group B-1.

Spore chain morphology: Umbellate Monoverticillate ( = Streptomyces Mature spore chains

Color of colony: Aer ia l mass c o l a i n the Red co lor -ser ies (grayish

Color i n medium: Melanoid pigments are not formed in peptone-yeast-

Reports f r o m


Figure 50. A. chromofuscus. Spi ra l spore chains. ( X 400) on oatmeal

F igure 51. A . chromofuscus. Spiny spores ; electron micrograph f rom

Figure 52. Streptoverticil l ium cinnamoneum. K V (umbellate monoverti-

F igure 53. Streptoverticil l ium cinnamoneum. Smooth spores ; electron

agar , 18 days .”

14 day culture on oatmeal agar . 31

cil late) spore chains (X 1000) on oatmeal agar, 14 days.

micrograph f r o m 14 day culture on yeas t -mal t a g a r .


rharnnose o r raffinose. P o o r growth on a l l carbon sources including - D- glucose may inaicate a requi rement f o r a growth factor not present in the basa l medium,

Actinomyces citreof luorescens Korenyako,' Krasil'nikov, Nikitina and Sokolova. Description: Korenyako e t at. 1960, 133-159. Type strain: INMI 2292 m.). INMI 2292.

chains long, often with m o r e than 50 spores p e r chain (Fig. 54). morphology i s seen on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 55).

yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar .

g ray ish yellow to light olive brown) on yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar .

yeast-iron agar , tyrosine a g a r o r tryptone-yeast broth. Yellow pigment i s found i n medium in glycerol-asparagine a g a r and yellow o r green pigment i s found in medium i n yeast-malt agar , oatmeal a g a r and sa l t s - s t a r c h a g a r . This pigment is not a pH indicator.

- D-fructose and rhamnose a r e utilized f o r growth. t r a c e of growth on sucrose, 1-inositol o r raffinose.

ISP 5265 f r o m N.A. Krasil 'nikov as USSR-RIA 648 = ISP description by Group B-2.

Spore chain morphology: Section Rectiflexibiles. Mature spore This

Color of colony: Aer ia l m a s s color in the Yellow color -ser ies on

Reverse side of colony: No distinctive pigments (light yellow o r

Color i n medium: Melanoid pigments a r e not produced in peptone-

Carbon utilization: _D-Glucose, &-arabinose, _D-xylose, _D-mannitol, No growth o r only

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 Cur t i s 1916, 110 (&ee Kutzner and Waksman 1959, 528-538). Type strain: CBS-R. MII'ller s t ra in (C.A. deVries, CBS personal communication). ISP 5233 f r o m G.A. deVr ies as CBS-R MUller strain.

Spore chain morphology: Section Rectiflexibiles. Mature spore chains usually 10 to 50, o r often m o r e than 50, spores p e r chain. (Fig. 56). s t a r c h a g a r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 57).

s t r a t e conidia on oatmeal a g a r , Emerson ' s potato dextrose a g a r and potato c a r r o t agar . Sclerotia formation on yeast-malt a g a r and oatmeal a g a r was repor ted by one observer .

Color i n colony: A e r i a l m a s s color in the Yellow color -ser ies on yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

Reverse side of colony: Light olive brown to strong brown o r very d a r k brown on yeast-malt a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r ; brown subs t ra te mycelium pigment on oatmeal a g a r is pH sensit ive changing f r o m brown to green with addition of 0.05 N NaOH. Potato plug i s pigmented d a r k blue in 4-7 days.


This morphology i s seen on yeast-malt agar , oatmeal a g a r , sa l t s -

Special morphological charac te r i s t ics : One observer repor t s sub-


Color in medium: Melanoid pigments a r e not formed in peptone- yeast-iron agar , tyrosine a g a r o r tryptone-yeast broth. Reddish, yellowish brown , o r greenish pigments a r e found i n the medium in yeast-matt a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and g lycero l -aspar - agine a g a r ; yellowish brown pigment is changed to green with addition of 0.05 N NaOH. greenish brown in 4-7 days.

and _D-fructose are utilized f o r growth. growth on sucrose , L-inositol, rhamnose and raffinose.

Liquid surrounding d a r k blue potato plug is pigmented

Carbon utilization: _D-Clucose, I=-arabinose, Q-xylose, _D-mannitol No growth o r only t r a c e of

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: 12531/54 (Ibrd. ; s e e a l s o Gauze Gottlieb 1968). Streptomyces coeruleorubidus (Preobrazhenskaya) Pr idham e t a l . 1958, 67. ISP 5145 f r o m T. P. Preobrazhenskaya a s 12531/54. ISP description by Group A-13.

bes 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 yeas t -mal t a g a r , oatmeal a g a r and glycerol-asparagine a g a r (Fig. 58b). Mature spore chains generally 10 to 50 s p o r e s p e r chain. Spore surface: Spiny (Fig. 59).

Color of colony: Aer ia l m a s s color in the Blue c o l o r - s e r i e s on oatmeat a g a r and sa l t s - s ta rch 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 agar .

mal t a g a r ; grayed yellow to yellowish green 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 green and grayed r e d o r orange on glycerol-asparagine a g a r . These subs t ra te mycelium pigments a r e not pH indicators when tested with 0.05 N NaOH o r HCl .

i ron agar , tyrosine a g a r and tryptone-yeast broth. found in medium in yeast-malt a g a r , sa l t s - s ta rch a g a r and glycerol- aspaxagine a g a r a f t e r 14 days ; i t i s not pH sensit ive.

Carbon utilization: _D-G lucose, &-arabino se, sucrose , _D -xy lose, - i.-inositol, 2- mannitol, _D-fmctose, rhamnose and raffinose a r e utilized for growth.

Spore chain morphology: Section Spirales. Sa l t s - s ta rch a g a r shows

Reverse side of colony: Grayish yellow to yellowish brown on yeas t -

Color in medium: Melanoid pigments a r e formed in peptone-yeast- Red pigment may be

Streptomyces cora lus Dietz (unpublished data). Description: Not published (Alma Dietz, Upjohn Go., personal communication, Sept. 1964). tion). ISP description by Group B -2.

Spore chain morphology: Section Rectiflexibiles (Fig. 60). Some long spore chains f o r m open loops suggesting morphology. Mature spore chains a r e usually very long, often with m o r e than 50 spores p e r chain. This morphology i s seen on yeast-malt a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 61).

co lor -ser ies on sa l t s - s ta rch 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 . Aer ia l mycelium is poorly developed on oatmeal a g a r .

Type s t ra in : UC 2098 = NRRL 2442 (Dietz, personal communica- ISP 5256 f r o m A. Dietz, Upjohn Go. as UC 2098 = NRRL 2442.

Color of colony: Mature a e r i a l m a s s color is usually in the Red

Figure 54. _A. citreofluorescens. R F spore chains (X 500) on oatmeal

F igure 55. A. citreofluorescens. Smooth spores ; electron micrograph

F i g u r e 56. S . coelicolor. RF spore chains (X 380) on Waksman's s t a r c h

F i g u r e 57. 2. coelicolor. Smooth spores ; electron micrograph f r o m

Figure 58. A. coeruleorubidus. Spiral spore chains (X 500): (a) Coils

- agar , 7 days.8

f r o m 21-day culture on glycerol-asparagine agar . 2 9

agar B.=* - 7 day culture on yeas t -mal t agar . 2 9

of many t u r n s on sa l t s - s ta rch agar, 24 days; (b) Spi ra l s on oatmeal agar, 24 d a y s . '' 18 day culture on oatmeal agar .

F igure 59. A. coeruleorubidus. Spiny spores ; electron micrograph f r o m


Reverse side of colony: P a l e o r light grayish yellow to orange- yellow on yeas t -mal t agar , oa tmeal a g a r and glycerol-asparagine a g a r ; light yellow t o gray ish r e d on sa l t s - s ta rch a g a r . pigment is a pH indicator changing f r o m yellow CR yellowish brown t o r e d with addition of 0.05 N NaOH.

Color in medium: Melanoid pigments a r e formed in peptone-yeast- i r o n a g a r , tyrosine a g a r and tryptone-yeast broth. found in 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 in the medium i n yeast-malt a g a r and oatmeal a g a r . is changed f r o m yellow to pink with the addition of 0.05 N NaOH.

- i-inositol, _D-mannitol, _D-fructose, rhamnose and raffinose a r e a l l utilized for growth.

Substrate mycelium

Red pigment is

This pigment

Carbon utilization: _D-Glucose, _L-arabinose, sucrose , 2-xylose,

k t i n o m y c e s cyanoalbus Krasil 'nikov and Agre. Description: Krasil 'nikov and Agre 1960, 273-274. Type s t r a i n INMX-414 (ibld., Krasil 'nikov personal communication). ISP 5198 f r o m N.A. KraSil 'nikov as INMI-414 = RIA 662.

Spore chain morphology: Section Spi ra les o r Rectiflexibiles. Distinct s p i r a l s are r a r e . Moderately shor 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 tu rns (Fig. 62). This morphology is s e e n on yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and 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 days) i s in the G r e e n c o l o r - s e r i e s on yeas t -mat t a g a r and sa l t s - s ta rch a g a r ; Green, Gray 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 . Aer ia l mycelium is usually poorly developed on oa tmeal a g a r .

Reverse s ide of colony: Grayed yellow to grayed yellow-green on yeas t -mal t a g a r and oa tmeal a g a r ; g rayed yellow-green to blue on s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . Subs t ra te mycelium pigment is not a pH indicator.

Color i n medium: Melanoid pigments are not formed in peptone- yeast-iron a g a r , ty ros ine a g a r o r tryptone-yeast broth. No pigment is found in the medium i n yeas t -mal t agar , oatmeal a g a r , sa l t s - s ta rch a g a r o r glycerol-aeparagine a g a r .

Carbon Utilization: _D-Glucose, _L-arabinose, sucrose , _D-xylose, - i-inoeitol, _D-mannitol, _D-fructose and rhamnose a r e utilized for growth. No growth o r only t r a c e of growth on raffinose.

ISP Description by Group A-13.

Streptomyces eh imens is Nakazawa. Description: Japanese Pa ten t 2125, Feb . 4, 1954; Japanese Pa ten t 8148, Nov. LO, 1955 (see a l s o Shibata, Honjo, Tokui and Nakazawa 1954, 168 f o r publication of name and antibiotic production only). Type strain: 138 = IF0 3398 (Nakazawa, personal communication; Shibata e t at. 1954, 168). Nakazawa, Takeda Chem. Ind., Ltd. as 138 = LF'O 3398. ISP description by Group B-9.

is usually umbellate-monoverticillate (biverticillate), but sporulating a e r i a l mycel ium i e v e r y sparBe o r absent on yeaat-malt agar, oatmeal a g a r , sa Lts - e tarch a g a r and glyce rot-aeparagiae a g a r . B ive rt icil late epore chains can b e seen on P r i d h a m and Gottlieb carbon utilization medium plus _D-xylose or _L-arabinose and somet imes can be found on oatmeal a g a r a f t e r 21 daye (Fig. 64). Spore chains are short , usually

ISP 5253 f r o m K.

Spore chain morphology: Section Verticil lati . Mature a e r i a l mycelium

Figure 60. S. coralus. E, very long spore chains (X 500) on oatmeal

F igure 61. S . coralus. Smooth spores ; electron micrograph f r o m 21

Figure 62. _A. cyanoalbus. (a) Open sp i ra l s and imperfect sp i ra l s

agar, 7-days. '

day c u l G r e on oatmeal agar . 2 9

(X 500) on glycerol-asparagine agar , 21 days. (b) Flexuous chains on yeas t -mal t agar , 21 days. 23

18 day culture on oatmeal agar .

(X 1100) on oatmeal agar , 21 days.

10 day culture on yeast-malt agar .

F igure 6 3 . A. cyanoalbus. Spiny spores ; electron micrograph f r o m

Figure 64. 2. ehimensis. Z V (umbellate monoverticillate) morphology

Figure 65. S, ehimensis. Smooth spores ; electron micrograph f r o m


with 3 to 10 s p o r e s p e r chain. Color of colony: A e r i a l m a s s color for m a t u r e sporulating a e r i a l

mycelium cannot be determined accura te ly because of poor sporulation on yeas t -mal t a g a r , sa t t s - s ta rch 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 is i n the G r a y co lor -ser ies . (The spore sur face of poorly sporulating growth is generally in the Yellow o r Red c o l o r - s e r i e s on yeas t -mal 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.)

Reverse side of colony: Brown t o d a r k brown on yeas t -mal t a g a r ; g ray ish yellow, orange yellow o r yellowish brown on oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . pigment is not a pH indicator.

Color in medium: Melanoid pigments a r e produced i n peptone-yeast- i ron 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 gray ish yellowish brown pigment may be found i n the medium i n yeas t -mal t a g a r , oa tmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . This pigment is not pH sensitive.

Carbon utilization: _D-Glucose, &inositol, _D-Mannitol and _D-fructose are utilized for growth. control and on other ISP carbon sources .

Spore surface: Smooth (Fig. 65).

Substrate mycelium

T r a c e s of growth occur on the carbon f r e e

S t rep tomyces endue Anderson and Cottl ieb 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. Patent 2 ,746,902; Upjohn Go. 1954, B r i t i s h Pa ten t Spec. 705,622. See also: Anderson and Gottlieb 1952, 294-308. Type strain: 9-20 = NRRL 2339 ( E d . ) . ISP 5187 f r o m D. Gottlieb as 9-20 = NRRL 2339.

(Fig. 66) . Individual s p o r e s are 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 Gray c o l o r - s e r i e s a f t e r 21 days on yeast-malt a g a r , oa tmeal 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 on glycerol-asparagine a g a r and young a e r i a l mycelium on o ther media m a y be white. Older colonies somet imes become mois t and black.

Reverse side of colony: No distinctive pigments (grayed yellow to yellowish brown o r gray) on yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s t a r c h a g a r and glycerol-asparagine a g a r . strate mycelium sometime s becomes mois t and black. Subs t ra te color is not affected by pH change.

i r o n agar , tyroeine a g a r o r tryptone-yeast broth. No pigments a r e found i n the medium i n yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r or glycerol-asparagine a g a r .

- D-fructose and rhamnose are utilized for growth. No growth o r only trace of growth on sucrose, &-inositol and raffinose.

ISP description by Group A-13. Spore chain morphology: Section Spi ra les with tightly closed s p i r a l s

In o lder cu l tures the oub-

Color i n medium: Melanoid pigments not formed i n peptone-yeast-

Carbon utilization: _D-Glucoee, _L-arabinose, _D-xylose, _D-mannitol,

Streptomyces fi l ipinensis Ammann, Gottlieb, Brock, G a r t e r and Whitfield. Description: Ammann e t at . 1955, 559-563. Type strain: 114-8 @d.). ISP 5112 f r o m D. Gottlieb as NRRL 2437 = 114-8. description by Group B-3.

ISP

F i g u r e 66. 2. endus. Spi ra l spore chains; tight s p i r a l s (X 550) on

F igure 67. S. e-s. Warty spores in tight sp i ra l s ; electron micrograph

Figure 68. S. filipinensis. Spi ra l spore chains. (a) Tight sp i ra l s a t end

-- glycerol-asparagine agar , 23 days. l5

f r o m 13-day culture on Gauze's medium No. 1.

of s p o r e c h a i n on yeas t -mal t agar , 15 days; (b) Open coils (X 500) on yeas t -mal t agar , 11 days. 2o

14 day culture on yeas t -mal t agar . lo

(X 625);n s a l t s - s t a r c h agar, 14 days.22 (b) I r regular sp i ra l s on yeast-malt agar , 21 days. 23

Figure 69. S. filipinensis. Spiny spores ; electron micrograph f r o m 1

Figure 70. A. finlayi. (a) Flexuous chains, hooks and imperfect sp i ra l s

F igure 71. _A. finlayi. Hai ry spores .2z


Spore chain morphology: Section Spirales. Tight s p i r a l s (Fig, 68a) o r open coils, usually of severa l tu rns (Fig. 68b), occur at the end of moderately tong spore chains of LO to 50 o r m o r e s p o r e s p e r chain on yeas t -mal t a g a r , oatmeal a g a r and sa l t s - s ta rch a g a r . Spi ra l s may be replaced by morphology on glycerol-asparagine a g a r . Spore surface: Spiny (Fig. 69).

yeast-matt a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine agar .

Reverse side of colony: No distinctive pigments (colorless to grayish yellow) on yeast-matt a g a r , oatmeal agar , sa l t s - s ta rch a g a r and g 1 y c e r o 1 -a spa r agine aga r . i ron a g a r , tyrosine a g a r and peptone-yeast broth. Pigments other than melanoids a r e not found in the medium i n yeas t -mal t a g a r , oatmeal agar , 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 r a g i n e a g a r .

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 and raffinose a r e utilized for growth. No growth o r only t r a c e of growth on rhamnose.

Color of colony: A e r i a l m a s s color is in the Gray c o l o r - s e r i e s on

Color in medium: Melanoid pigments a r e produced in peptone-yeast-

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 = FBUA 1869. Group B-12.

Spore chain morphology: Section Rectiflexibiles to Spirales. Short spore 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 imperfec t s p i r a l s of only one o r two turns (Fig. 70a); longer chains may f o r m open s p i r a l s of th ree o r four tu rns (Fig. 70b). Hooks and loops on s h o r t spore chains a r e not charac te r i s t ic of morphology. 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 ; s p i r a l s a r e bes t developed on yeas t -mal t a g a r and oatmeal a g a r . Spore surface: Hairy (Fig. 71).

mal t agar , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

g reen (grayish yellow to olive brown on yeas t -mal t a g a r and oatmeal a g a r ; pale greenish yellow o r yellow-green to moderate o r d a r k olive green on sa l t s - s ta rch a g a r o r glycerol-asparagine agar ) . Substrate mycelium pigment is not a pH indicator.

Color in medium: Melanoid pigments are not produced i n peptone- yeas t - i ron agar , tyrosine a g a r o r tryptone-yeast broth. No pigments a r e found in the medium in yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

growth a l s o occurs on _L-arabinose, _D-xylose and rhamnose, although growth on these carbon sources is Less than on _D- glucose. of sucrose is doubtful. - D-mannitol, _D-fructose and raffinose.

ISP description by

Color of colony: Aer ia l m a s s color in the Gray c o l o r - s e r i e s on yeas t -

Reverse side of colony: Grayed yellow to yellow-brown modified by

Carbon utilization: _D-Glucose is utilized for growth. Significant

Utilization No growth o r only t r a c e of growth oni - inos i to l ,

Actinomyces f luorescens Korenyako, Krasil'nikov, Nikitina and Sokolova. Description: Korenyako e t a l . 1960, 133-159 (see a l s o Engl. Trans . , 128-156). Type strain: INMI 592 e.). ISP 5203 f r o m N.A. Krasil 'nikov a s INMI 592 = RIA 647. ISP Description by Group B-6.


Spore chain morphology: Section Rectiflexibiles (Fig. 72). Mature spore chains generally LO to 50 spores p e r chain. This morphology is seen on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine a g a r . Spore surface: Smooth (Fig. 73).

yeas t -mal t agar , oatmeai a g a r and sa l t s - s ta rch a g a r ; White o r Yellow color - s e r i e s on glycerol-asparagine agar .

Reverse side of colony: No distinctive pigments (colorless to grayish yellow on oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r ; grayed orange-yellow to olive brown on yeas t -mal t agar). Reverse pigment i s not pH sensit ive o r is changed only slightly f r o m yellow-brown to yellow-green, with addition of 0.05 N NaOH.

yeas t - i ron agar , tyrosine a g a r o r tryptone-yeast broth. A smal l amount of yellow pigment may be found in the medium in yeast-malt agar , oat- m e a l a g a r , sa l t s - s ta rch a g a r and tryptone-yeast broth. This pigment when present is 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 for growth. t r a c e of growth on sucrose , L-inositol o r raffinose. cription f o r t h i s species (=. c.) s ta tes that rhamnose i s not utilized , noting that this i s one difference b e t w e e n k . f luorescens and other spec ies in the fluorescent group. recorded good growth on rhamnose, equivalent to growth on _D-grlucose.

Color of colony: Aer ia l m a s s color i n the Yellow color -ser ies on

Color in medium: Melanoid pigments a r e not produced in peptone-

No growth or only The original des-

However, a l l of the 3 ISP o b s e r v e r s

Streptomyces f rag i l i s Anderson, Ehrlich, Sun and Burkholder. Des- cription: Anderson e t a l . 1956, 100-105 and 114-115. Type strain: isolate C 1437 = P-D 04926 = NRRL 2424 w.), Res . Lab., Parke , Davis and Co. a s P - D 04926 = NRRL 2424. ISP description by Group B-2.

Spore chain morphology: Section Spirales to Rectiflexibiles. Short spore chains i n many straight to flexuous chains (Fig. 74a), but hooks o r loops of s m a l l d iameter and incomplete sp i ra l s a r e a l s o common (Fig. 74b). Spi ra l s of two o r m o r e complete turns a r e a l s o present, but a r e not common, Spore chains are generally short, but usually contain LO o r m o r e spores p e r chain. tendency toward warty sur face on spores near the te rmina l p a r t of the chain (Fig. 75).

mal t agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . and g lycerol-asparagine agar .

Reverse side of colony: Grayed yellow to yellow-brown modified by r e d (Light to moderate yellow o r orange-yellow on sa l t s - s ta rch a g a r ; light gray ish reddish brown to dark brown on yeast-malt agar , oatmeal a g a r and glycerol-asparagine agar). to yellow brown with addition of 0.05 'N HC1.

yeast-iron agar , tyrosine a g a r or trypto ne-yeast broth. Medium may remain unpigmented o r may contain smal l amounts of yellow o r red- brown pigment in the mdeium in yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r or glycerol-asparagine agar . This pigment, when present,

ISP 5044 f r o m L. Anderson,

Spore surface: Smooth with some

Color of colony: A e r i a l m a s s color i n the Red co lor -ser ies on yeas t -

White a e r i a l mycelium may a l s o be present on sa l t s - s ta rch a g a r

Reddish brown pigment may change


F i g u r e 72 . A. fluorescens. RF spore chains (X 500) on s a l t s - s t a r c h

F igure 73. S. f luorescens. Smooth spores ; electron micrograph f r o m

Figure 74. S. fragil is . (X 380) on Waksman's s t a r c h a g a r B, 21 days:

- agar , 23 days. l5

21 day culture on yeast-malt agar . 32

(a) Straight t o flexuous chains; (b) Loops, incomplete s p i r a l s and shor t sp i ra l s on the s a m e medium,'*

F igure 75. S. fragil is . Smooth spores , with some sur face i r regular i t ies on te rmina l spores ; electron micrograph f r o m 14 day culture on yeas t -mal t a g a r . 2 9

-


is pH sensitive changing f r o m red-brown to yellow with addition of 0.05 N HC1.

utilized for growth. only t r a c e of growth o n 2 -inositol, _D-mannitol, 2 - f ruc tose , rhamnose o r raffinose.

Carbon utilization: D_-Glucose, _L-arabinose and 2-xylose a r e Utilization of sucrose is doubtful. No growth o r

Actinomyces fulvoviridis Kuchaeva, Krasillnikov, Skryabin and Taptykova. Description: Kuchaeva e t a l . 1960, 251 (see a l s o Engl. t rans l . , 222-25L). Type strain: INMI 16-3 (i&. ; type s t ra in selected by N.A. Krasillnikov, personal communication). Krasil 'nikov a s INMI 16-3 = RIA 660.

spore chains moderately shor t with LO to 50 spores p e r chain. This morphology is seen on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 77).

yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

Reverse side of colony: Yellow to yellow-brown plus t r a c e s of green (pale yellow, yellow o r grayed greenish yellow to olive brown o r dark olive on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine agar ) .

yeast-iron agar , 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 in medium i n yeast-malt agar , oatmeal agar , sa l t s - s ta rch 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 for growth; utilization of sucrose 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.

ISP 5210 f r o m N.A. ISP description by Group B-12.


Color of colony: Aer ia l m a s s color i n the Gray co lor -ser ies on

Reverse pigment is not a pH indicator. Color in medium: Melanoid pigments a r e not produced in peptone -

Streptomyces ga lbus F r o m m e r . Description: Fromme r 1959, 187- 206. 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 , personal communication). ISP 5089 f r o m C. W. Hesselt ine as 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. Sp i ra l s may be open coils of severa l turns, tight sp i ra l s a t the ends of flexuous spore chains o r imperfec t sp i ra l s and loops a t the ends of spore chains suggesting =morphology (Fig. 78). Spore chains mod- e ra te ly long with LO to 50, o r often m o r e than 50, spores p e r chain. Spore surface: Smooth to warty. Surface i r regular i t ies suggesting very smal l war t s a r e charac te r i s t ic (Fig. 79).

yeast-malt a g a r and oatmeal a g a r and in the Yellow color -ser ies o r Gray co lor -ser ies on sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . Gray color i s represented by tabs 2fe (medium gray) o r 3fe (light brown- i s h gray) on a l l media; and Iba o r 2ba (pale yel1ow)or 1 1/2db (pale gray- i s h yellow) on sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

brown o r olive brown) on yeast-malt agar , oatmeal agar , sa l t s - s ta rch agar and glycerol-asparagine agar .


Reverse side of colony: No distinctive pigments (yellow to yellow-

Figure 76. A. fulvoviridis. RF spore chains on yeast-malt agar,

Figure 77. A. fulvoviridis. Smooth spores; electron micrograph from

Figure 78. S. galbus. Spiral o r R A spore chains; (a) On oatmeal agar ,

Figure 7 9 . S. galbus. Smooth spores with surface irregularit ies

21 days.23

14 day culture on salts-starch agar."

14 day<(X 400). (b) On glycerol-asparagine agar, 16 days. 33

suggesting very small warts; electron micrograph from 16 day culture on yeast-malt agar. '

- -


Color in medium: Melanoid pigments a r e produced in peptone-yeast- i r o n a g a r and tryptone-yeast broth, but not in tyrosine agar . Yellow soluble pigment i s found in the medium i n yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and gtycero 1-asparagine agar . This pigment i s not a pH indicator.

Carbon utilization: ,D- Glucose, _L-arabinose, _D-xylose, j--inositol, - D-mannitol and _D-fructose are utilized for growth. Only a t r a c e of growth comparable to growth on the carbon-free control is found on sucrose, rhamnose and raffinose.

Streptomyces gardner i (Waksman 1942) Waksman 1961. Descriptions: Proactinomyces 2. Gardner and Chain 1942, 123-127; Proactinomyces gardner 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 gardner i (Waksman) Waksman 1961, 215-216. Type strain: IMRU 3834 (Waksman 1961, 216). as IMRU 3834.

chains moderately long, usually LO to 50, o r often m o r e than 50, spores p e r chain. This morphology is seen on yeast-malt agar , oatmeal agar , sa t t s - s ta rch a g a r and glycerol-asparagine a g a r , a tthough a e r i a l mycelium may be poorly developed on oatmeal a g a r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 81).

sa l t s - s ta rch a g a r in 14 to 21 days. inadequate for determination of a e r i a l m a s s color on yeast-malt agar , oatmeal a g a r and glycerol-asparagine agar . good sporulating a e r i a l growth i n the Gray co lor -ser ies on _ . potato a g a r (potato 200 gm; a g a r 18 gm; tap water 1 L; pH 6.9-7.1).

yellow o r grayed yellow-brown) on yeast-malt a g a r , oatmeal agar , sa l t s - s t a r c h agar , glycerol-asparagine agar and potato agar .

i ron a g a r and tryptone-yeast broth, but not i n tyrosine agar (or only in t r a c e amounts) in 2 to 4 days. No pigments a r e found in the medium in yeast-malt a g a r , oatmeal agar , sa l t s - s ta rch a g a r o r glycerot-asparagine a g a r .

Carbon Utilization: _D-GLucose, _L-arabinose, sucrose, ,D-xylose, - D-fructose, rhamnose and raffinose a r e utilized for growth. o r only t r a c e of growth oni - inos i to l and _D-mannitol. :

ISP 5064 f r o m S.A. Waksman ISP description by Group B -3.

Spore chain morphology: Section Rectiftexibiles (Fig. 80). Spore

Color of colony: Aer ia l m a s s color in the G r a y co lor -ser ies on Mature a e r i a l mycelium may be

One observer obtained

Reverse side of colony: No distinctive pigment (colorless to grayish

Color in medium: Melanoid pigments a r e produced in peptone-yeast-

No growth

Streptomyces getaticus (Waksman) Waksman and Henrici. Descrip- tion: Actinomyces Waksman 1919, 165-166; Streptomyces ge la t icus (Waksman) Waksman and Henrici 1948, 952 (see a l s o Anderson g s. 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 descr ip- tion by Group B -6.

mycelium is not procuded on most media. Absence of a e r i a l mycelium on most media was a l s o noted in the original description (Waksman 1919 3. c&.) which, however, r e c o r d s open sp i ra l s on the scant growth on synthetic agar . ISP o b s e r v e r s found wavy spore chains, but no sp i ra l s i n the thin a e r i a l mycelium on yeast-matt a g a r and gtycerol-asparagine

Spore chain morphology: Section Rectiflexibiles (Fig. 82). Aer ia l

3 2 1 I N T E R N A T I O N A L J O U R N A L

a g a r . The absence of sp i ra l s has a l s o been noted by Anderson e t a l . (1956%. &.), by Waksman in a later description (1961o~.&t.), and by HUtter ( 1 9 6 7 9 . A t . ) . Spore surface: Smooth (Fig. 83) .

When a n adequate spore m a s s is produced i t is in the G r a y co lor -ser ies . This was observed on yeast-malt a g a r and 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 agar , Lemco a g a r and milk a g a r .

yellow) on yeas t -mal t a g a r , oatmeat agar , sa l t s - s ta rch a g a r o r glycerol- asparagine a g a r .

yeas t - i ron a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment is found in medium in yeas t -mal t agar, oatmeal a g a r , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

Carbon utilization: Q-Glucose, sucrose , _D-xylose and rhamnose a r e utilized for growth. - i'-inositol, Qmanni to l , Q-fructose and raffinose.

Color of colony: Aer ia l mycel ium i s poorly developed on most media.

Reverse side of colony: No distinctive pigments (colorless to grayish

Color i n medium: Melanoid pigments a r e not produced in peptone-

No growth o r only t r a c e of growth on I=-arabinose,

Actinomyce s g lobisporus Kra si llnikov. Description: K r a si llnikov 1941, 48. Streptomyces globisporus (Krasil 'nikov) Waksman 1953, 39. Type s t ra in : INMI 2302 = RLA 335 ( N A . Krasil 'nikov, personal 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 spore 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 yeas t -mal 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 lycero l -aspar - agine a g a r . Spore surface: Smooth (Fig. 85).

yeas t -mal 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 lycero l -aspar - agine a g a r .

Reverse side of colony: No distinctive pigments (pale yellow to light olive brown) on yeas t -mal t a g a r , oatmeal agar , 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 in peptone- yeast-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 yeas t -mal 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 sensit ive when tested with 0.05 N HC1 o r NaOH.

- D-fructose and rhamnoie a r e utilized for growth, No grou?h o r only t r a c e of growth on sucrose , i-inositol o r raffinose.

Color of colony: Aer ia l m a s s color in the Yellow c o l o r - s e r i e s on

Carbon utilization: D-Glucose, L-arabinose, D-xylose, D-mannitol,

- Streptomyces goshikiensis Niida. Description: Niida, Japanese

Pa ten t 425909, Japan Official Gazette 38-26948 (Engl. Trans l . f r o m T. Niida, personal communication). Streptomyces goshik iemis Niida Kondo, Sakamoto and Yumoto 1961, 365 and in 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 sp i ra l s of 4 to

Name without description:

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

Figure 81. S. gardneri. Smooth spores; electron micrograph from

Figure 82. 2. gelaticus. Figure 83. 2. elaticus. Smooth spores, Au/Pd shadowed; electron

Figure 84. _A. globisporus. RF spore chains on: (a) Oatmeal agar, 7 (b) WakGan’s starch agar B, 21 days (X 380).”

Figure 85. 2. globisporus. Smooth spores; electron micrograph from

16 -

14 day &tu=yeast-malt agar. lo

micrograp! from 21 day culture on oatxheal agar.

days (X 1000).

14 day culture on oatmeal agar. 2 9

spore chains on salts-starch agar, 23 days.”


10 convolutions a t the end of long spore chains (Fig. 86). chains LO to 50 s p o r e s p e r chain; longer chains are often observed. This morphology i s seen on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . Spore surface: Smooth with minor s u r f a c e i r regular i t ies , but no t rue spines (Fig. 87).

on yeas t -mal t agar , oatmeal a g a r and sa l t s - s ta rch agar . color on these media is between 3ge (light gray ish yellowish brown) and 5dc o r 6ec (grayish yellowish pink) color tabs of Tresner -Backus Color Wheela. Aeria l mycelium may be poorly developed on glycerol-asparagine a g a r .

Reverse side of colony: No distinctive pigments (charac te r i s t ic grayed yellow on oatmeal agar , 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 yeas t -mal t agar ) .

C o l a in medium: Melanoid pigments a r e formed in peptone-yeast- i r o n agar , tyrosine a g a r and tryptone-yeast broth. P igments other than melanoids o r faint t r a c e s of yellow are not produced i n yeast-malt a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

for growth. f r e e basa l medium on L-arabinose, sucrose , _D-xylose, L-inositol, - D-rnannitol, rhamnose and raffinose.

Mature spore

Color of colony: Aer ia l m a s s color i n the Red (or Gray) c o l o r - s e r i e s Charac te r i s t ic

Carbon utilization: _D-Glucose and possibly _D-fructose are utilized Only a t r a c e of growth comparable to growth on carbon-

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 gr i se inus Waksman 1959, 1043-1046. Type strain: IMRU 3478 (Waksman 1959, 1045). 3478.

Spore chain morphology: Section Rectiflexibles (Fig. 88). Mature spore chains predominantly flexuous and 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 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. 89).

Color of colony: Aer ia l mass color i n the Yellow c o l o r - s e r i e s on sa l t s - s ta rch a g a r and glycerol-asparagine a g a r and i n the Yellow o r Gray c o l o r - s e r i e s on oatmeal a g a r and yeast-malt agar . Neares t matching color tab f r o m the Gray co lor -ser ies is Zdc, yellowish gray.

yellowish brown or light olive brown) on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar . pigment may be changed slightly f r o m pale yellow to pale yellow-brown o r pale violet with addition of 0 .05 N NaOH, or f r o m yellowish brown to pale yellowwith 0 .05 N HC1.

ISP 5047 f r o m S.A. Waksman as IMRU ISP description by Group B-3.

Reverse side of colony: No distinctive pigments (grayish yetlow to

The subs t ra te mycelium

Color in medium: Melanoid pigments a r e not produced i n peptone- yeas t - i ron agar , ty ros ine a g a r o r tryptone-yeast broth. yellow to reddish brown pigment may be found i n the medium i n oatmeal agar . f r o m pale yellow to pale violet with addition of 0 . 0 5 N NaOH.

- D-fructose and rhamnose are utilized for growth. t r a c e of growth on sucrose , 1-inositol or raffinose.

A t r a c e of

This pigment, when preeent, is slightly pH sensit ive changing

Carbon utilization: _D-Glucose, _L-arabinose, _D-xylose, _D-mandtol, No growth o r only

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,

Figure 87. 2. goshikiensis. Smooth spores with minor surface irregu- 22

larities, but no true spines. chromium shadowed. Electron micrograph from 22 day culture on oatmeal agar. *'

Figure 88. 2. griseinus. Figure 89. 2. grieeinus. Smooth spores; electron micrograph from

spore chains on yeast-malt agar, 15 days. l1

14 day culture on oatmeal agar. lo


ISP 5191 f r o m M. Oda, Meiji Seika Co. as 1068. Group A-13.

Spore chain morphology: Section Spi ra les (Fig. 90). Mature spore chains generally LO to 50 s p o r e s p e r chain. This morphology i s observed on yeas t -mal t a g a r , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-aspar - agine agar . Spore surface: Smooth (Fig. 91).

on yeast-malt a g a r , oa tmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine agar . Charac te r i s t ic color is between 4ge o r 5ge (light grayish reddish brown) and 4ig (tight gray ish brown) color tabs of Tresner -Backus color wheels.

Reverse side of colony: No distinctive pigments on yeas t -mal 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 .

Color i n medium: Melanoid pigments not formed in peptone-yeast- i r o n agar , tyrosine a g a r o r tryptone-yeast broth. medium i n yeas t -mal t a g a r , oa tmeal agar , sa l t s - s ta rch a g a r o r glycerol- asparagine agar .

and _D-fructose are utilized for growth. growth on sucrose , i- inositol , rhamnose and raffinose.

ISP description by

Color of colony: Aer ia l m a s s color in the Red o r G r a y c o l o r - s e r i e s

No pigment found in

Carbon utilization: _D-Glucose, &-arabinose, _D-xylose, 2 -manni to l No growth o r only t r a c e of

Ac tinomyc e s g ri s e oincarnatu s Pr eob ra zhe nskaya , Rya bova and BLinov. Description: Preobrazhenskaya e t at. &Gauze et at. 1957, 169- 171. Type strain: 9673/55 (C. F. Gauze, personal communication: G. F. Gauze, cited by Gottlieb 1968, 20) . Streptomyces griseoincarnatus (Preobrazhenskaya e t aL.) P r i d h a m et at. 1958, 69. ISP 5274 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 Spi ra l s a r e generally open with 5 to 7 o r m o r e turns (Fig. 92a);flexuous spore chains and imperfec t s p i r a l s suggesting &morphology are a l s o common (Fig. 92b). Spi ra l s a r e bes t developed on oatmeal a g a r and sa l t s - s ta rch a g a r . Spore chains a r e moderately long, especially on oatmeal a g a r and sa l t s - s ta rch a g a r with LO to 50, o r often m o r e than 50, s p o r e s p e r chain. Spore s u r - face $piny (Fig. 93).

Color of colony: Aer ia l m a s s color in the Gray c o l o r - s e r i e s on sa l t s - s t a r c h a g a r a f t e r 14-21 days ; younger a e r i a l mycelium may be in the Red co lor -ser ies . Aer ia l m a s s color in the Gray o r B e d c o l o r - s e r i e s on yeas t -mal 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 represent ing the a e r i a l color, but a l l chose tabs ranging f r o m grayish yellowish pink or grayish reddish brown to medium gray.

yellow on ea l t s - s ta rch a g a r ; orange-yellow to reddish brown on yeas t - mal t a g a r , oatmeal a g a r and glycerol-asparagine agar ) , mycelium pigment is not a pH indicator.

yeas t - i ron agar , tyrosine a g a r o r tryptone-yeast broth. Red o r orange pigment is found in the medium i n yeast-malt a g a r i n 7 days and in oatmeal a g a r in 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,

mannitol, _D’-fructose and rhamnose a r e utilized for growth. Reports vary on utilization of i - inos i to l and only a t r a c e of growth is found on raffinose.

Reverse side of colony: Yellow t o yellow-brown plus r e d (grayish

Substrate


Carbon utilization: _D-Glucose, &-arabinose, sucrose , ,D -xylose, _D-

Figure 90. 2. griseofuscus. S iral spore chains (X 500) on glycerol-

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 salts- starch agar, 14 days. “

Figure 93. A. griseoincarnatus. Spiny spores; electron micrograph from 14day culture on oatmeal agar. 22

asparagine agar, 22 days. +

3 30 INT ERNATIONA L J O U R N A L

Actinomyces griaeoruber Ryabova and Preobrazhenskaya. Descrip- tion: Ryabova and Preobrazhenskaya in Gauze e t at. 1957, 186 and 195. Streptomyces Rriseoruber (Ryabova and Preobrazhenskaya) Pridham -- et 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, 20) . 2022/55.

moderately long with LO to 50, o r often more than 50, spores per chain. This morphology is seen on yeast-malt agar, oatmeal agar, salts-starch agar and glycerol-asparagine agar. Spore surface: Spiny (Fig. 95).

malt agar, oatmeal agar o r salts-starch agar. One observer placed aer ia l m a s s color in both the Gray and Red color-series on these media, Good ae r i a l na ss growth is not found on glycerol-asparagine agar. Color tabs selected by observers indicate a range from light o r medium gray to yellowish o r brownish gray in the Gray series, to grayish yellowish pink in the Red ser ies ; gray i s the predominant color.

orange to grayish red o r grayish reddish brown) on yeast-malt agar, oatmeal agar, salts-starch agar and glycerol-asparagine agar. The substrate mycelium pigment is not a pH indicator.

yeast-iron agar, tyrosine agar o r tryptone-yeast broth. not found in the medium in yeast-malt agar, oatmeal agar or salts- starch agar. A t race of yellow pigment is found in glycerol-asparagine agar after 14-21 days; i t is not pH aensitive.

- i-inositol, _D-mannitol, _D-fructose, rhamnose and raffinose a r e a l l utilized for growth.

ISP 5275 from C.F. Gauze as

Spore chain morphology: Section Spirales (Fig. 94). Spore chains ISP description by Group B-5.

Color of colony: Aerial mass color in the Gray color-series on yeast-

Reverse side of colony: Yellow to yellow-brown plus red (moderate

Color in medium: Melanoid pigments a r e not produced in peptone- Pigments a r e

Carbon utilization: ,D -Glucose, 5-arabinose, sucrose, _D-xylose,

Streptomycee Rriseoruber Y amaguchi and Saburi. Description: Yamaguchi and Saburi 1955, 203 and 219-231. Not Streptomyces griseoruber (Ryabova and Preobrazhenskaya) Pridham et at. 1958, 69. strain: H-4650 E d . ) . ISP 5281 from T. Yamaguchi as H-4650. ISP description by Group B-LO.

Spore chain morphology: Section Spirales o r Retinaculiaperti. Open o r tight spirals a r e produced a t the ends of moderately long spore chains of LO to 50 o r more spores per chain (Fig. 96a). complete, forming only 1 o r 2 turns o r a hook at the end of the spore chain. Straight to flexuous chains without hooks o r spirals a r e a lso present (Fig. 96b). agine agar, but may a l so be observed on yeast-malt agar, oatmeal agar and ealts-starch agar. Spore surface: Smooth (Fig. 9 7 ) .

on yeast-malt agar, oatmeal agar, salts-starch aga r and glycerol- aeparagine agar. reddish brown) from the Gray color-series and tab 5ge (also light grayish reddish brown) from the Red color-serieB.

Reverse aide of colony: Yellow to yellow-brown plus red (reddish orange o r reddieh brown on yeast-malt agar, oatmeal agar and salts- s tarch agar; tight yellowish pink on glycerol-asparagine agar). Sub- s t ra te mycelium pigment is a pH indicator changing from reddish orange

Type

Spirals a r e often in-

This morphology is usually best on glycerol-aspar-

Color of colony: Aerial mass color is in the Red o r Gray color-series

The color is represented by tab 5fe (light grayish

Figure 94. A. griseoruber Ryabova and Preobrazhenskaya. Spiral spore

Figure 95. A. griseoruber Ryabova and Preobrazhenskaya. Spiny spores;

Figure 96. S. griseoruber Yamaguchi and Saburi. Spiral or RA spore

chains @ 400) on yeast-malt agar, 14 days. ’ electronmicrograph from 16 day culture on yeast-malt agar.’

chains on glycerol-asparagine atar, 14 days: (a) Open a n x i g h t spirals a t ends of long chains (X 580); (b) Straight to glexmua chains and some terminal hooks or spirals (X 230). l6

electronmicrograph from 21 day culture on yeast-malt agar. l7 Figure 97. S. griseoruber Yamaguchi and Saburi. Smooth spores;


to purple with addition of 0.05 N NaOH and from reddish orange to yellowish orange with addition of 0.05 N HC1.

i ron agar, tyrosine agar and tryptone-yeast broth. ment (depending upon pH) may be found in the medium in yeast-malt agar , oatmeal agar, salts-starch agar and glycerol-asparagine agar. This pigment is pH sensitive, changing from orange to purple with addition of 0.05 N NaOH and from orange t6 yellow with addition.of 0 . 0 5 N HC1.

- D-fructose and rhamnose a r e utilized for growth. No growth o r only t race of growth on sucrose, _D-mannitol and raffinose.

Color i n medium: Melanoid pigments a r e produced in peptone-yeast- Yellow to red pig-

Carbon uti lization : _D -G luco s e , _L -a ra bin0 s e , ,D - xy lo s e , i - i no si to 1,

Streptomyces Kriseoviridis Anderson, Ehrlich, Sun and Burkholder . Description: Streptomyces griseoviridus (91c) Anderson e t at. 1956, 109-115. (Spelling changed to 5. griseoviridis by Waksman 1961, 225, and by L. E. Anderson, personal communication, Sept., 1967. ) Type strain: A 9071 = P-D 04955 = NRRL 2427 (%). Anderson as P - D 04955 = NRRL 2427, IMRU 3735. Group B -6.

Spore chain morphology: Section Spirales (Fig. 98). Some open spirals have only one o r two turns o r a r e poorly developed suggesting - F U morphology. Spore chains moderately long with LO to 50 spores per chain. Spore surface: Smooth (Fig. 99).

Color of colony: Aerial mass color in the Red color-series (grayish yellowish pink to pale orange yellow) on yeast-malt agar , oatmeal agar, salts-starch agar and glycerol-asparagine agar. t h i s culture in the Yellow color-series. (Anderson G. , 1956) included the following statement: "Aerial mycelium is white to pink-tan, pink-gray, o r brown o r occasionally greenish". The greenish color w a s not observed on ISP media.

on yeast-malt agar, oatmeal agar, salts-starch agar and glycerol- asparagine agar . is changed only slightly by 0.05 N NaOH o r HCl.

i ron agar, tyrosine agar o r tryptone-yeast broth. No pigment in medium, o r only t race of yellow, in yeast-malt agar, oatmeal agar , sal ts-s tarch aga r o r glycerol-asparagine agar.

and rhamnose a r e utilized for growth. doubtful. raffinos e.

ISP 5229 from L.E. ISP description by

One observer placed The original description

Reverse side of colony: Grayed yellow to olive brown o r dark brown

Substrate mycelium pigment is not a pH indicator o r

Color in medium: Melanoid pigments a r e not found in peptone-yeast-

Carbon utilization: D, -G luco se, &-arabinose, _D-xy lo s e, _D -mannito 1 Utilization of _D-fructose is

No growth o r only t race of growth on sucrose, i- inositol o r

Streptomyces gr iseus (Krainsky) Waksman and Henrici. Descriptions: Actinomyces griseus Krainsky 1914, 682; Actinomyces griseus (Krainsky) Wakeman 1916, 119 and 128; Waksman 1919, 123-125; Strepto- myces gr iseus (Krainsky) Waksman and Henrici 1948, 948. (See also Actinomyces globieporus va r .s t reptomy cini Kra si l'nikov 1949, LOO ; Xctinomyees etreptomycini Krasil'nikov 1957, 598). IMRU 3463 tneotype, Waksman 1961, 226). as IMRU 3463.

Type strain: ISP 5236 from S.A. Waksman

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

Figure 99. S. griseoviridis. Smooth spores, Au/Pd shadowed; electron

Figure 100. S. griseus.

Figure 101. 2. griseus. Smooth spores; electron micrograph from

agar, 22 days. Is

micrograph from 21 day culture on oatmeal agar. yI

14 days.

14 day culture on yeast-malt agar. l6

spore chains (X 230) on yeast-malt agar,


spore chains moderately long with LO to 50 s p o r e s p e r chain. morphology is seen on yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 101 ).

(2db, pale yellow, o r 1 1/2 db - 1 1/2 ec, pale greenish yellow) on yeas t - mal t 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 .

Reverse side of colony: No distinctive pigments (grayed yellow to olive brown o r light yellowish brown) on yeast-malt a g a r , oa tmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

but not i n peptone-yeast-iron a g a r o r tryptone-yeast broth. other than t r a c e s of yellow a r e found i n the medium in yeas t -mal t agar , oatmeal a g a r , sa l t s - s ta rch a g a r o r glycerol-asparagine agar .

Carbon utilization: Q-Glucose, _D-xylose, _D-mannitol and _D-fructose a r e utilized for growth. No growth o r only t r a c e of growth on &-arabinose, sucrose , i- inositol , rhamnose o r raffinose.

This

Color of colony: Aer ia l m a s s color i n the Yellow color -ser ies

Color in medium: Melanoid pigments a r e formed in tyrosine a g a r , No pigments

Streptomyces herbar ico lor Kawato and Shinobu. Description: Kawato and Shinobu 1959, 114-119. Type strain: 608 (ibid.) ISP 5123 f r o m R. Shinobu as 608.

spore chains generally LO to 50 s p o r e s p e r chain on yeas t -mal t a g a r and glycerol-asparagine a g a r . developed o r absent on oatmeal a g a r and s a l t s - s t a r c h a g a r . surface: Smooth (Fig. 103).

glycerol-asparagine a g a r and yeas t -mat t a g a r . Aer ia l mycelium is poorly developed on a l l ISP media; mass color cannot be determined on oatmeal a g a r or sa l t s - s ta rch a g a r .

to olive brown or brown o r i t may contain a n additional pH sensit ive pigment with a color range f r o m pale purple o r pa le purplish pink to green o r greenish blue. When this pigment is present, addition of 0.05 N NaOH changes color f r o m violet-red to green o r blue and this change is r e v e r s e d by addition of 0.05 N HC1. pigment was found by one observer i n yeast-malt agar , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine agar .

Color i n medium: Melanoid pigments are not formed o r occur only in t r a c e amounts in peptone-yeast-iron a g a r and tyrosine agar . Pig- ment may o r may not be formed in the medium i n yeas t -mal 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 rans ien t subs t ra te pigment.

Carbon utilization: 2-Glucose, &-arabinose, sucrose , _D-xylose, - D-fructose and raffinose a r e utilized for growth. No growth o r only t r a c e of growth on i- inositol , _D-rnannitol and rhamnose.

ISP description by Group B-12. Spore chain morphology: Section Rectiflexibiles (Fig. 102). Mature

Sporutating a e r i a l mycelium is poorly Spore

Color of colony: Aer ia l m a s s color i n the Gray c o l o r - s e r i e s on

Reverse side of colony: Substrate mycelium may be grayish yellow

The pH sensit ive mycelial

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. Patents 2,931,756 and 2,950, 277. production only, without description: Nakazawa and Shibata & Imamura -- e t at . 1956, 648-653.

Name and antibiotic

Type strain: 23572 (Nakazawa e t a t , , H.).

Figure 102. S . herbaricolor. g epore chains (X 625) on yeast-malt

Figure 103. S . herbaricolor. Smooth spores; electron micrograph from

Figure 104. S . humidus. Spiral spore chains (X 600) on oatmeal agar,

Figure 105. S. humidus. Smooth spores; electron micrograph from

Figure 106. S. indigoferus. Portion of colony (X 400) showing R F spore

agar, 14-day 8 . '' 14 day c z t u r e on yeast-malt agar . 2Z

14 days.76 - 2 1 day c z h x a t m e a l agar. *' chains a t edge of colony (top) and coremia on colony s u r f a c e (bottom).

14 day culture on yeast-malt agar. 30

Figure 107. 2. indigoferus. Smooth spores; electron micrograph from


ISP 5263 f r o m E. Lesse l , Amer . Type Cult. Cot. as ATCC 12760 f r o m Takeda Pharm. Ind., LTD. as 23572 = P O 3520. Group B -10.

s p i r a l s resembling morphology (Fig. 104). Spore chains shor t to moderately long with LO to 50 s p o r e s p e r chain. This morphology i s seen on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine a g a r , Spore surface: Smooth (Fig. 105).

tion repor t s production of black, moist (hygroscopic) a r e a s on mature a e r i a l mycelium, this was not recorded on ISP repor t s .

s e r i e s (3fe o r 31i, brownish gray, f r o m the Gray c o l o r - s e r i e s to 5dc o r 5cb, gray ish yellowish pink, in the Red co lor -ser ies ) .

grayish yellow o r tight olive gray) 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 .

Color i n medium: Me tanoid pigments not produced in peptone-yeast- i ron 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, is found in the medium i n yeas t -mal t agar , oatmeal a g a r , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

- D-mannitol, _D-fructose and rhamnose a r e utilized for growth. growth o r only t r a c e of growth on s u c r o s e and raffinose.

ISP description by

Spore chain morphology: Section Spirales with many incomplete

Special morphological charac te r i s t ics : Although the original descr ip-

Color of colony: Aer ia l m a s s color in the Gray o r Red color-

Reverse side of colony: No distinctive pigments (colorless, pale o r

Carbon utilization: 9-Glucose, k a r a b i a o s e , _D-xylose, i- inositol , No

Streptomyces indigoferus Shinobu and Kawato. Descriptions: Shinobu and Kawato 1960b, 49-33. Type strain: 709 @bid. ). ISP 5124 f r o m R. Shinobu as 709. ISP description by Group B-5.

chains moderately long with LO to 5 0 , o r somet imes m o r e than 50, s p o r e s p e r chain. Aer ia l mycelium is poorly developed on a l l ISP media and is obscured by extensive coremia formation on yeas t -mat t agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r (Fig. 106). Spore s u r - face: Smooth (Fig. 107).

Color of colony: Aer ia 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 coremia formation on yeas t -mal t a g a r , sa l t s - s ta rch a g a r aad glycerol-asparagine a g a r . When adequate a e r i a l mycelium is formed, i t is 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 . glycerol- u r e a a g a r and soil ex t rac t a g a r by one observer .

less, grayish yellow o r d a r k brown on yeas t -mal t agar , oa tmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar . A t rans ien t blue to r e d pigment (depending on pH) may a l s o b e found i n the mycelium on these media. This pigment changes f r o m blue o r co lor less to r e d with addition of 0.05 N HC1.

Color i n medium: Melanoid pigments a r e formed i n peptone-yeast- i ron a g a r and tryptone-yeast broth, but a r e not formed in tyrosine agar . P igments o ther than melanoids were not found i n the medium in yeas t - m a t t agar , oatmeal a g a r , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

Carbon utilization: _D-Glucose and _L-arabinose are utilized for

Spore chain morphology: Section Rectiflexibiles. Mature spore

Aer ia l mass color i n the Gray c o l o r - s e r i e s was a l s o found on

Reverse side of colony: Substrate mycelium is usually nearly co lor -


growth. No growth o r only t r a c e of growth on sucrose, i- inositol , Q-mannitol, _D-fructose, rhamnose or raffinose.

Utilization of _D-xylose is doubtful.

Actinomyces janthinus Artamonova and Krasillnikov. Description: Artamonova and Krasil 'nikov 1960, 334 (see a l s o English translation, 3-7 and 300-306). Type strain: 117 (w); type s t ra in selected by N.A. Krasil'nikov, personal communication. Krasillnikov a s INMI 117 = RIA 659. ISP description by Group B-8.

moderately long with LO to 50, or sometimes m o r e than 50, spores p e r chain. This morphology is seen on yeast-malt agar , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine agar . Spore surface: Spiny (Fig. 109).

on yeas t -mal t a g a r and sa l t s - s ta rch a g a r ; White co lor -ser ies on oatmeal a g a r and White o r Gray co lor -ser ies on glycerol-asparagine agar . neares t color tabs in the Red co lor -ser ies a r e 5dc, 5ca, 5cb and 6ec (all grayish yellowish pink).

yellow or pale yellow) on yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r or glycerol-asparagine agar .

Color i n medium: Melanoid pigments a r e formed i n peptone-yeast- i ron a g a r and tryptone-yeast broth, but not in tyrosine agar . pigment is found in the medium in sa l t s - s ta rch a g a r . This pigment i s not pH sensit ive when tested with 0.05 N NaOH or HCI.

i -inositol, _D-mannitol, _D-fructose, rhamnose and raffinose a r e a l l utilized for growth.

ISP 5206 f r o m N-A.

Spore chain morphology: Section Spi ra les (Fig. LOB). Spore chains

Color of colony: Aer ia l m a s s color i n the Red o r White co lor -ser ies

The

Reverse side of colony: No distinctive pigments (colorless to grayed

Yellow

Carbon utilization: 2-Glucose, &-arabinose, sucrose, _D-xylose,

Str eptoverticillium (SLC) (Barr and Carman) Baldacci, Farina and Locci. Descriptions: Streptomyces kentuchensis B a r r and Ca r m a n 195 6, 2 86 - 2 89 ; St reptovertici 1 lium kentuchense B a lda cci, Farina and Locci, 1966, 161. Type strain: 5x1 RCV a s Streptomyces kentuckensis B a r r and Carman (ibid). ISP 5052 f r o m F.S. B a r r a s ATCC 12691 = 5x1 RCV.

Section Verticil lati , biverticil late) (Fig. 110). moderately long with LO to 50 spores p e r chain.

ISP description by Group B-1. Spore chain morphology: Umbellate Monoverticillate (= Streptomyces

Mature spore chains This morphology i s

seen on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine agar . Spore surface: Smooth (Fig. 111).

Color of colony: Aer ia l m a s s color i n the Red co lor -ser ies on yeas t - mal t agar , oatmeal a g a r and sa l t s - s ta rch a g a r ; Red or White color- s e r i e s on glycerol-asparagine agar .

Reverse side of colony: No distinct pigments (grayed yellow on oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r to brown on yeas t -ma tt agar ) .

Color i n medium: Melanoid pigments a r e produced i n peptone-yeast- i r o n agar and tryptone-yeast broth, but.not in tyrosine agar . No pigment is found in the medium i n yeast-matt agar , oatmeal agar , sa l t s - s t a r c h a g a r or glycerol-asparagine agar .

Carbon utilization: _D-Glucose, L-inositol and fructose a r e utilized for 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 (umbellate- monoverticillate) 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. '


of growth on L_-arabinose, sucrose, _D-mannitol, rhamnose and raffinose.

Actinomyces lavendocolor Kuchaeva, Krasil'nikov, Taptikova and Gesheva. Description: Kuchaeva et at. 1962, 103-124. Type strain: INA 4518 (ibid.). 749.

chains a r e long, often with m o r e than 50, spores p e r chain (Fig. 112). This morphology is seen on yeast-malt a g a r , oatmeal agar , sa l t s - s t a r c h a g a r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 113).

mal t agar , oatmeal a g a r and sa l t s - s ta rch agar . A e r i a l mycelium is poorly developed on glycerol-asparagine agar .

Reverse side of colony: No distinct pigments (nearly co lor less to pale yellow o r very pale yellowish pink) on yeast-malt a g a r , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar .

Color in medium: Melanoid pigments a r e formed in peptone-yeast- i r o n a g a r and usually i n tyrosine a g a r and tryptone-yeast broth. P igments other than melanoids a r e not found i n the medium i n yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

inositol are utilized for growth. tot, _D-fructose, rhamnose and raffinose resembles growth on the carbon f r e e control.

ISP 5216 f r o m N.A. Krasil 'nikov as INA 4518 = RIA ISP description by Group B-12.

Spore chain morphology: Section Retinaculiaperti o r E. Spore

Color of colony: Aer ia l m a s s color i n the Red co lor -ser ies on yeast-

Carbon utilization: _D-Glucose, &-arabinose, 2-xylose, a n d i - T r a c e of growth on sucrose, _D-manni-

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 chains often terminate in sp i ra l s o r hooks and loops of wide d iameter (Fig. 114). than 50, spores p e r chain. agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 115).

Color of colony: Aer ia l m a s s color i n the Red co lor -ser ies (Grayish yellowish pink o r yellowish pink) on yeast-malt agar , oatmeal a g a r and sa l t s - s ta rch agar .

Reverse side of colony: No distinctive pigment (pale yellow to light yellowish brown) on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol asparagine agar .

Color i n medium: Melanoid pigments are formed in peptone-yeast- i r o n a g a r and usually in tyrosine a g a r and tryptone-yeast broth. P igments other than melanoids are not formed i n the medium in yeast-malt a g a r , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine agar .

inositol a r e utilized for growth. No growth o r only trace of growth on sucrose , e m a n n i t o l , rhamnose o r raffinose.

Spore chain morphology: Section Retinaculiaperti I Long, f Lexuous

Mature spore chains have LO to 50, o r often m o r e This morphology is seen on yeast-malt

Carbon utilization: _D-Glucose, &-arabinose, _D-xylose and i- Utilization of f ruc tose is doubtful.

Figure 112. A. lavendocolor. Very long flexuous chains of more than 50 spores ( 5 4 8 5 ) on salts-starch agar. 37 Loops and hooks, representative of F U cultures a r e also reported for this strain.

14 day culture on salts-starch agar. "

loops onGatmea1 agar , 21 days: (a) (X 300); (b) (X 1200).23

14 day culture on salts-starch agar . "

Figure 113. 4. lavendocolor. Smooth spores; electron micrograph from

Figure 114. A. lavendofoliae. R A spore chains with terminal spirals or

Figure 115. A. lavendofoliae. Smooth spores: electron micrograph from


Actinomyces levoris Nikitina. Descriptions: Nikitina 1958, 142; Korenyako, Krasillnikov and Nikitina 1960, 116-L32 (see a l so Engl. Transl. , 111-127). Type strain: INMI 2725 (Ibld.; type s t ra in selected by N.A. Krasil'nikov, personal communication). Krasillnikov a s INMI 2725 = RIA 338. ISP description by Group B-5.

chains generally long with LO to 50, o r often more than 50, spores p e r chain (Fig. 116). agar , sa l t s - s ta rch aga r and glycerol-asparagine agar . SmoQth (Eig. 117).

mal t agar , sa l t s - s ta rch aga r and glycerol-asparagine agar . Aerial mycelium may be inadequate fo r color observation on oatmeal agar .

grayish greenish yellow to orange-yellow o r yellowish brown) on yeast- mal t agar , oatmeal agar , sa l t s - s ta rch aga r and glycerol-asparagine agar .

Color in medium: Melanoid pigments a r e not formed in peptone- yeast-iron agar , tyrosine aga r o r tryptone-yeast broth. found in the medium i n yeast-malt agar, oatmeal agar , sa l t s - s ta rch agar o r glycerol-asparagine aga r .

and 2- f ruc tose a r e utilized for growth. growth on sucrose, i--inositol, rhamnose and raffinose.

ISP 5202 f r o m N.A.

Spore chain morphology: Section Rectiflexibiles. Mature spore

This morphology is seen on yeast-malt agar , oatmeal Spore surface:

Color of colony: Aer ia l m a s s color i n Yellow color -ser ies on yeast-

Reverse side of colony: No distinctive pigment (Grayish yellow o r

No pigment is

Carbon uti liza tion: ,D -G luco s e , _L-a rabino s e, ,D -xyLo s e, _D -mannit01 No growth o r only t race of

Streptomyces lilacinus Nakazawa, Tanabe, Shibata, Miyake and Takewaka. Descriptions: Nakazawa e t at. 1956, 81; Nakazawa e t at. 1957, 111-115. as IF0 3944 = 2305.

cillate (biverticillate), (Fig. 118). 3-10 spores p e r chain. oatmeal agar , sa l t s - s ta rch aga r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 119).

yellowish pink) on yeast-malt agar , oatmeal agar , salts-starch a g a r and g lyc e r o 1 -a spa rag ine aga r .

Reverse side of colony: Yellow-brown plus r ed (grayish reddish brown to strong brown) on yeast-malt agar , oatmeal agar , sa l t s - s ta rch aga r and glycerol-asparagine aga r ; this pigment is somewhat pH sensitive changing f r o m brown to reddish brown with addition of 0.05 N NaOH and f rom brown to yellowish brown with addition of 0.05 N HC1.

i ron agar , tyrosine aga r and tryptone-yeast broth. found in the medium in yeast-malt agar, oatmeal agar , sa l t s - s ta rch aga r and glycerol-asparagine agar . tested with 0.05 N NaOH o r HCl, showing the same change noted fo r subs t ra te color.

on positive utilization ofi-inositol . on _L-arabinose, sucrose, g-xylose, _D-mann.itol, _D-fructose, rhamnose and raffinose.

Type strain: 2305 u ) F I S P 5254 f r o m K. Nakazawa

Spore chain morphology: Section VerticiLlati, umbellate monoverti- Spore chains a r e short, usually with

ISP description by Group B-LO.

This morphology is seen on yeast-malt agar ,

Color of colony: Aer ia l m a s s color i n the Red co lor -ser ies ( grayish

Color in medium: Melanoid pigments a r e formed in peptone-yeast- Red pigment is

This pigment is pH sensit ive when

Carbon utilization: 2-Glucose is utilized fo r growth; repor t s vary No growth o r only t r ace of growth


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 a i w 4417 56 (neotype, Preobrashenskaya, personal communication: N.A. Krasil 'nikov, personal communication, January, 1968). ISP 5166 f r o m T.P. P reobrazhenskaya as 4417/56. Group B-1.

chains moderately short , usually LO to 50 spores p e r chain. surface: Spiny (Fig. 121). (One observer found smooth spores . )

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 yeas t -mat t agar , oatmeal a g a r and glycerol-asparagine a g a r ; White o r Red (5cb grayish 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 .

Reverse side of colony: No distinctive pigments (light o r grayish yellow on oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r to olive brown o r brown on yeas t -mal t agar).

Color in medium: Melanoid pigments a r e produced i n peptone-yeast i ron a g a r , tyrosine a g a r and tryptone-yeast broth. P igments other than melanoids a r e not found in the medium in yeast-malt a g a r , oatmeal agar , sa l t s - s ta rch 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, rhamnose and raffinose a r e a l l utilized for growth.

ISP description by

Spore chain morphology: Section Spi ra les (Fig. 120). Mature spore Spore

Actinomyce s longi sporus -f lavus Kra sil'nikov. Descriptions : Krasil 'nikov 1941, 30. pe r s ona 1 communication ; K r a si 1' nikov, p e r s ona 1 co mm uni ca ti 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 as 5165.

Spore chain morphology: Section Spi ra les . Open s p i r a l s ; spore chains resembling morphology are a l s o common (Fig. 122). Mature spore 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. agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 123).

sa l t s - s ta rch a g a r . other ISP media.

yellow o r light gray ish yellowish brown) on yeas t -mal t a g a r , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

yeas t - i ron a g a r , tyrosine a g a r o r tryptone-yeast broth. found in medium i n yeas t -mal t agar , oatmeal a g a r , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

and r h a m n o e are utilized f o r growth. growth on s u c r o s e and raffinose. mannitol is doubtful.

Type strain: 8 1/53 (neotype, Preobrazhenskaya,

ISP description by Group A-9.

This morphology may be observed on yeas t -mal t

Color of colony: A e r i a l nass color is in Yellow color -ser ies on Aer ia l mycelium is poorly developed o r absent on

Reverse side of colony: No distinct pigments (colorless, grayish

Color i n medium: Melanoid pigments a r e not formed i n peptone- No pigment

Carbon utilization: _D-Clucose, L.-arabinose, _D-xylose, D - f r u c t o s e No growth o r only t r a c e of

Utilization of i - inos i to l and 2-

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

Figure 117. _A. levoris. Smooth spores; electron micrograph from 32

Figure 118. 2. lilacinus. BIV (umbellate monoverticllate) morphology

Figure 119. 2. lilacinus. Smooth spores; electron micrograph from

Figure 120. _A. lonfisporus. Spiral spore chains (X 640) on oatmeal

Figure 121. ,A. longisporus. Spiny spores; electron micrograph from

agar, 5 Z y s i -

day culture on salts- starch agar.

(X 400) on salts-starch agar, 14 days.”

21 day culture on oatmeal agar.’’

agar, 21 days.

14 day culture on salts-starch agar.’


S p o r e chain morphology: Section Verticil lati , umbellate monoverti- ci l late (biverticillate, F ig . 124). Mature spore 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 sa l t s - s ta rch a g a r or yeas t - matt a g a r . matt a g a r , oatmeat a g a r and glycerol-asparagine agar . Absence of good a e r i a l mycelium on var ious media was a l s o noted by Shinobu i n h is original description &, c&.). Spore surface: Smooth.

s e r i e s on sa l t s - s ta rch a g a r ; i t is absent o r poorly developed (White o r Yellow) on other ISP media.

Reverse side of colony: No distinctive pigments (light brown to grayed yellow) on yeas t -mat t a g a r , oatmeal agar , sa l t s - s ta rch a g a r o r g Lyce r o 1 -a sparagine agar .

i ron a g a r , tyrosine a g a r and tryptone-yeast broth. found in medium i n yeas t -mal t a g a r , oatmeal a g a r and sa l t s - s ta rch a g a r a f te r L4-21 days; it is not a pH indicator.

Carbon utilization: _D-Glucose, i- inositol a d D - f m t t o s e a r e utilized for growth. No growth o r only t r a c e of growth on &-arabinose, sucrose , _D-xylose, _D-mannitol, rhamnose and raffinose.

Y a m a s h i t a , Isono, Anzai, Serizawa and Sekiyama. Description: Sawazaki et at. 1955, 44-45. communication). S r o u p B-5.

Spore chain morphology. Section Verticil lati , umbellate monoverticil late (biverticillate, Fig. 125). LO, o r somet imes m o r e than LO, s p o r e s p e r chain. Spore surface: Smooth (Fig. 126).

yeas t -mal t a g a r , oatmeal agar , sa l t s - s ta rch a g a r and g lycero l -aspar - agine a g a r . glycerol-asparagine a g a r .

Reverse side of colony: No distinctive pigments (grayish yellow to orange yellow on oa tmeal a g a r , sa l t s - s ta rch a g a r and glycerol- asparagine a g a r ; olive brown to brown on yeas t -mal t agar ) .

i r o n a g a r and tryptone-yeast broth, but not i n ty ros ine a g a r . No pigment is found i n the medium i n yeas t -mal t a g a r , oatmeal agar , sa 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, sucrose and _D-fructose a r e utilized for growth. t r a c e of growth on &-arabinose, _D-xylose, _D-mannitol, rhamnose and raffinose.

Aer ia l mycelium is poorly developed or absent on yeas t -

Color of colony: Mature a e r i a l mycelium i n the Red o r Yellow color-

Color in medium: Melanoid pigments a r e formed in peptone-yeast- Yellow pigment i s

Streptomyces mashuens is Sawazaki, Susuki, Nakamura, Kawasaki,

Type strain: 449 (S. Suski, personal ISP 5221 f r o m S. Susuki as 449. ISP description by

Spore chains a r e shor t with 3 to

Color of colony: Aer ia l mass color i n the Red c o l o r - s e r i e s on

Mature aerial mycelium may be poorly developed on

Color in medium: Melanoid pigments are produced i n peptone-yeast-

Utilization of 1-inositol is doubtful. No growth o r only

Streptomyces matens is Margalith, Bere t ta and Timbal. Description: Margalith et at. 1959, 71-75. Type s t ra in : ME/17 k d , ) . ISP 5188 f r o m L.G. Silvestri , Lepetit S.p.A. as Lep. M-ME/l7. tion by Group A-13.

dominant f o r m s (Fig. 127); spore chains suggesting primitive s p i r a l s of cultures o r flexuous chains of =are a l s o present. Mature spore chains moderately long with LO to 50 o r m o r e spores p e r chain.

ISP descr ip-

Spore chain morphology: Section Spirales. Open s p i r a l s a r e the

Figure 122. A. longisporus-flaws. (a) Spiral spore chains (X 400) on (b) Terminal spirals and hooks on yeast- oatmealagar, 14 days.

malt agar, 14 days. '' Figure 123. A. longisporus-flaws. Smooth spores; electron micro-

Figure 124. S. luteoverticillatus. BIV (umbellate monoverticillate)

Figure 125. S. mashuensis. - BIV (umbellate monoverticillate) morphology

Figure 126. S. mashuensis. Smooth spores; electron micrograph from

graph from 21 day culture on salts-starch agar. '' morphology on yea st -malt agar?

(X 400) oatmeal agar, 14 days. ' 10 day c z t u r e on yeast-malt agar.'


This morphology is seen on yeast-malt agar, oatmeal agar, salts- starch agar and glycerol-asparagine agar. Spore surface: Spiny. Spines a r e short and inconspicuous (Fig. 128); some smooth spores may be seen.

(color tab 3ge, tight grayish yellowish brown; to 5fe, light grayish reddish brown) on salts-starch agar; Gray or White color-series on yeast-malt agar and glycerol-asparagine agar. One observer reports tab 5dc (grayish yellowish pink) from the Red color-series for growth on yeast-malt agar, oatmeal agar and salts-starch agar. This trace of red (or violet) pigment in aerial color is also noted in the original description (Margalith et at. 1959).

terist ic substrate colors to dark reddish brown on yeast-malt agar and salts-starch agar, tight grayish brown or yellowish pink on oatmeal agar and glycerol-asparagine agar. i s not a strong pH indicator. to pink with addition of 0.05 N HC1. )

iron agar, tyrosine agar or tryptone-yeast broth. a r e found in yeast-malt agar, oatmeal agar and glycerol-asparagine a g a r . 0.05 N HCl.

- D-mannitol, _D-fructose and rhamnose a r e utilized for growth. growth or only trace of growth on raffinose. doubtful.

Color of colony: Mature aerial m a s s color in the Gray color-series

Reverse side of colony: Red pigments in mycelium modify charac-

The substrate mycelium pigment (One observer reports change from violet

Color in medium: Melanoid pigments a r e not found in peptone-yeast- Red or violet pigments

One observer reports a change from violet to red by addition of

Carbon utilization: 2-Glucose, &-arabinose, _D-xylose, i-inositol,

Utilization of sucrose i s No

Streptomycea mediocidicus Okami, Utahara, Nakamura and Umezawa. Description: Okami et at. 1954, 98-100. Type strain: 535-A1 (ibid.). ISP 5021 from Y. Okami a s 535-A1 = MHJ 266. Group B -2.

on any ISP media or on 15 additional media tested by cooperators. Single conidia were observed on the substrate mycelium on glucose- asparagine agar and Waksman's starch agar B (Fig. 129). crystals formed on the substrate growth on yeast-malt agar (Fig. 130) . The original description (Okami et at. 1954) indicates that aerial mycelium w a s only poorly developed or absent on most media. not observed.

of the media tried by the cooperators.

agar, oatmeal agar, salts-starch agar and glycerol-asparagine agar. Thie pigment is not pH sensitive.

Color in medium: Melanoid pigments a r e formed in peptone-yeast- iron agar and tryptone-yeast broth, but may not be formed in tyrosine agar. Trace of yellow pigment may be found in the medium in yeast- malt agar, oatmeal agar, salts-starch agar and glycerol-asparagine agar; this pigment is not pH sensitive.

Carbon utilization: D_-Glucose, &-inositol and _D-fructose a r e utilized for growth. No growth or only trace of growth on &-arabinose, sucrose, - D-xylose, _D-mannitol, rhamnose and raffinose.

ISP description by

Spore chain morphology: Sporulating aerial mycelium is not produced

Masses of

Spore surface:

Color of colony: No sporulating aerial mycelium was found on any

Reverse side of colony: Yellow to greenish yellow on yeast-malt

Figure 127. S. matensis. Spiral spore chains on yeast-malt agar. Figure 128. 2. matensis. Spiny spores; spines a r e short and sparse.

Figure 129. S. mediocidicus. Single spores on substrate mycelium:

-- - Electron micrograph from 13 day culture on Gauze medium No. 1.’

(a) On Waksmanls starch agar B, 21 days.te (b) On glucoee- asparagine agar, 10 days (X 1000). *

Figure 130. S. mediocidicus. Crystalline m a m a on surface of eub- stratal growth (X 380) on yeast-malt agar, 21 days.

3 48 IN 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 from Y. Yagisawa f rom T. Hata, Kitasato Inst. a s KITA V-1179. ISP description by Group B-2.

Spore chain morphology: Section Rectiflexibiles (Fig. 131). Mature spore chains long with LO to 50, o r often more than 50, spores per chain. This morphology is seen on yeast-malt agar , oatmeal agar, sal ts-s tarch agar and glycerol-asparagine agar . Spore surface: Smooth (Fig. 132).

grayish yellowish pink) on oatmeal agar and salts-starch agar; White o r Gray color-ser ies on yeast-malt aga r and salts-starch agar . Mature sporulating aer ia l mycelium may be poorly developed on yeast- matt agar and salts-starch agar .

Reverse side of colony: Grayed yellow to yellow-brown is usually modified by red (becoming dark brown to reddish brown) on yeast-malt agar, oatmeat agar, salts-starch agar and glycerol-asparagine aga r ; t h i s pigment is not a pH indicator.

Color in medium: Melanoid pigments a r e formed in peptone-yeast- i ron agar, tyrosine agar and tryptone-yeast broth. Yellow o r red pigment is found in the medium in yeast-malt agar , oatmeal agar, sal ts- s tarch agar and glycerol-asparagine agar. sensitive when tested with 0.05 N NaOH o r HC1.

- D-mannitol, 2-fructose and raffinose a r e utilized for growth. vary on utilization of sucrose. rhamnose.

Color of colony: Aerial mass color in the Red color-ser ies (5cb,

This pigment is not pH

Carbon utilization: _D-Glucose, &-arabinose, D-xylose, i-inositol, Reports

No growth o r only t race of growth with

Streptomyces misakiensis Nakamura. Description: Nakamura 1961, 86-89; name and antibiotic production, without description, in Isono, Anzai and Suzuki 1958, 264-267. Type strain: 1755 (Nakamura, s.). ISP 5222 from S. Suouki, Inst. Phys. and Chem. Res., Tokyo a s 7617 = 1755.

spore chains generally tong with LO to 50, o r sometimes more than 50, spores pe r chain. meal agar, sal ts-s tarch aga r and glycerol-asparagine agar . Spore surfauz. Smooth (Fig. 134).

Color of colony: Aerial mass color in the Gray color-ser ies on salts-starch aga r and glycerol-aeparagine agar; Gray o r Red color- s e r i e s on yeast-malt aga r and oatmeal agar. color tab from the Gray color-wheel is 2dc (yellowish gray) for a l l ISP media. One observer selected tab 5dc (grayish yellowish pink) f rom the Red color-wheel as the most representative color on yeast- malt agar and oatmeal agar.

Reverse side of colony: Yellow to yellow-brown is modified by red, becoming brown to strong brown on yeast-malt agar, sal ts- starch agar and glycerol-asparagine aga r and orange to tight brown on oatmeal aga r ; t h i s pigment is not a pH indicator.

Color i n medium: Melanoid pigments a r e not formed in peptone- yeast-iron agar, tyrosine aga r o r tryptone-yeast broth. is found in the medium in yeast-matt agar, oatmeal aga r and salts- s tarch agar . NaOH or HCl.

ISP description by Group B-6. Spore chain morphology: Section Rectiflexibiles (Fig. 133). Mature

This morphology is seen on yeast-malt agar, oat-

The most representative

Red pigment

This pigment is not pH sensitive when tested with 0.05

Figure 131. S. melanogenes. R F spore chains. (a) (X 380), Waksman's

Figure 132. S. melanogenes. Smooth spores; electron micrograph from

Figure 133. S. misakiensis. R F spore chains (X 350) onyeadt-malt

Figure 134. S. misakiensis. Smooth spores; electron micrograph

starch agar B, 21 days.'' (b)(X 500), oatmeal agar, 7 days.'

21 day culture on oatmeal agar. 29

agar, 2 1-day s.

(Au/Pd axadowed) from 21 day culture on oatmeal agar.

-


Carbon utilization: _D-GLucose, sucrose , _D-fructose and raffinose a r e utilized f o r growth. arabinose, _D-xylose, i- inositol , _D-mannitol and rhamnose. original description (Nakamura, K. &. ) L_-arabinose and _D-mannitot were a l s o reported as supporting growth with Czapek's a g a r as basa l medium.

No growth o r only trace of growth with A- In the

Streptomyces naganishii Yamaguchi and Saburi. Description: Yamaguchi and Saburi 1955, 216-219 and 226. Type strain: H-4871

k d . ) . Group B -11.

Spore chain morphology: Section Spi ra les (Fig. 135a, b). Mature spore chains shor 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 spore chains on sa l t s - s ta rch a g a r and glycerol-asparagine a g a r may f o r m incomplete s p i r a l s o r hooks (Fig. 135c). oatmeal a g a r is especially poor. atypical club-Like swellings instead of s p o r e s (Fig. 136). Smooth (Fig. 137).

on yeast-matt a g a r and on carbon utilization medium plus _D-Glucose. The sporulated a e r i a l growth on oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r is usually white.

Reverse side of colony: P a l e o r grayish yellow t o yellow-brown modified in spots o r a t margins by r e d (dark pink, purplish pink o r gray ish reddish brown).

Color in medium: Melanoid pigments a r e formed in peptone-yeast- i ron 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 rans ien t r e d pigment is found i n the medium in 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 sensit ive when tes ted with 0.05 N NaOH or HC1.

Carbon utilization: _D-GLucose, _L-arabinose, _D-xylose, i- inositol , - D-mannitol, _D-fructose and rhamnose are utilized for growth. tion of raffinose is doubtful. sucrose .

ISP 5282 f r o m T. Yamaguchi as H-4871. ISP description by

Sporulation on Aer ia l hyphae m a y te rmina te i n

Spore surface:

Color of colony: Aer ia l m a s s color in the Gray o r Red c o l o r - s e r i e s

Subs t ra te pigment is not a pH indicator.

Utiliza- No growth o r only trace of growth with

Streptoverticil l ium ne t rops is (Finlay, Hochstein, Sobin and Murphy) Baldacci, Farina and Locci. Description: Streptomyces ne t rops is , U. S. Pa ten t 2,586,762, Feb. 19,1952; name and antibiotic production, without description, i n Finlay, Hochstein, Sobin and Murphy 1951, 341-343. 1966, 161-162. Type strain: Isolate 2937-6 = Pf izer F .D. 4779 = NRRL 2268. ISP 5259 f r o m J. Routien, Chas. P f i z e r and Go., Inc. as F. D. 4779 = NRRL 2268. ISP description by Group B-6.

Section Verticil lati , biverticillate). (BIV), flexuous, o r sp i ra l (BBV-2) spore chains (Fig. 138a, b, c). spore chains generally LO to 20 s p o r e s p e r chain. seen on yeas t -mat t a g a r , oatmeal agar , s a l t s - s t a r c h a g a r and glycerol- asparagine agar . Spore surface: Smooth (Fig. 139).

Color of colony: Aer ia l m a s s color i n the Red co lor -ser ies on yeas t - mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

Streptoverticil l ium netropsis, Baldacci, F a r i n a and Locci

Spore chain morphology: Umbellate monoverticil late (= Streptomyces Whorls may be composed of s t ra ight

Mature This morphology i s

Figure 135. 2. na anishii. a) S iral spore chains (X 200) on glycerol- asparagine ag:r, 14 day(s. *Spiral chains (X 400) on yeast-malt agar. ' 9 ( c ) Short chains with incomplete spirals and hooks, yeast- malt agar, 14 days.36

swellingc(%*ieast-malt agar. 36

14 day culture on glycerol-asparagine agar.

Figure 136. S. na anishii Aerial hyphae terminating in club-like

Figure 137. 2. naganishii. Smooth spores; electron micrograph from

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 microgra-ed) frzm 21 day culture on oatmeal agar.u

S Y S T E M A T I C B A C T E R I O L O G Y 3 5 3

Reverse side of colony: No distinctive pigments (grayish yellow to olive brown o r brown) on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerbl-asparagine agar .

Color in medium: Melanoid pigments are formed in peptone-yeast- i ron agar , but not in tyrosine agar . yellow o r brown pigment, is found in the medium in yeast-malt agar , oatmeal a g a r and sa l t s - s ta rch a g a r . slightly pH sensit ive changing f r o m pate yellow to pale pink when tested with 0.05 N NaOH.

for growth. sucrose , _D-xylose, _D-mannitol, rhamnose and raffinose.

No pigment o r only a t r a c e of

Pigment, if present, may be

Carbon utilization: _D-Glucose, L-inositol and _D-fructose a r e utilized No growth o r only t r a c e of growth wi th L_-arabinose,

Actinomyces n igrescens Sveshnikova. Description: Sveshnikova in Gauze e t at . 1957, 146-148. G.F. Gauze and regarded by him a s identical with Streptomyces hygro- scopicus, personal communication, Sept., 1964). Streptomyces nigre- scene (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.

chains moderately long with LO to 50, o r somet imes m o r e than 50, s p o r e s p e r chain. a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine agar . Smooth to warty (Fig. 141).

mal t agar , oatmeal agar , sa t t s - s ta rch 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 gray a e r i a l mycellium to a black shiny m a s s i n aging cultures. cooperators.

yellow) on yeast-malt a g a r , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine a g a r .

i r o n agar , tyrosine a g a r o r tryptone-yeast broth. No pigment is found in the medium in yeast-malt agar , oatmeal agar , sa t t s - s ta rch a g a r o r glycerol-asparagine a g a r .

D-mannitol, _D-fructose and raffinose a r e utilized for growth. or only t r a c e of growth with

Type strain: 1800/54 ( E d . ; selected by

ISP description by Group B -6. Spore chain morphology: Section Spirales (Fig. 140). Mature spore

This morphology is seen on yeast-malt agar , oatmeal Spore surface:

Color of colony: Aer ia l m a s s color in the Gray co lor -ser ies on yeas t -

This phenomenon was not observed by ISP

Reverse side of colony: No distinctive pigments (colorless to gray ish

Color i n medium: Melanoid pigments a r e not formed i n peptone-yeast-

Carbon utilization: ~-GLUCOSe, sucrose, _D-xylose, i- inositol , No growth

-arabinose and rhamnose.

Streptomyces nodosus Trejo. Description: Tre jo in Waksman 1961, 250; Tre jo and Bennett 1963, 436-439. Type strain: SC 2388 = ATCC .~

14899 (Trejo, personal communication). SC 2388.

chains may be poorly developed showing only a few turns o r hooks of smal l d iameter ; o r tightly knotted sp i ra l s may be formed (Fig. 142). Spore chains a r e short , often with only 3 to LO spores p e r chain. This morphology is seen on yeast-malt agar , sa l t s - s ta rch a g a r and glycerol- asparagine agar . Spore surface: Smooth, but some wrinkles o r folds a r e found on the spore surface (Fig. 143).

ISP 5109 f r o m W. H. Tre jo a s ISP description by Group A-12.

Spore chain morphology: Section Spirales. Spi ra l s on short spore

Figure 140. ,A. ni rescens

Figure 141. A. nigrescsns. Smooth spores (Au/Pd shadowed) with

Figure 142. 2. nodosus. Spirals on shqrt chains, some forming tight

Figure 143. 2. nodosus. Smooth spores; electron micrograph from

S i ra l spore chains (X 300) on salts-starch agar, 21 days. -%------ - small suFface irregularities suggesting warts.

"knots" (X 400); salts-starch agar, 21 days. '' 14 day culture on salts-starch agar. *'


Color of colony: Aer ia l m a s s color in the Gray co lor -ser ies on yeast-malt agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r ,

Reverse side of colony: Grayed yellow o r grayed greenish yellow to d a r k olive brown o r near black on yeast-malt agar , oatmeal agar , 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 is not a pH indicator.

i ron agar , tyrosine a g a r and tryptone-yeast ex t rac t broth. red or yellow orange pigment may be found in the medium in yeast-malt agar , oatmeal a g a r and glycerol-asparagine a g a r . this pigmert is pH sensitive changing f r o m r e d to green by addition of 0.05 N NaOH and f r o m green to blue-red by 0.05 N HC1.

Carbon uti t i za tion: _D -G Lucose, ,D - x y l o se, i - i n o sito 1, _D -mannito 1, - D-fructose and rhamnose a r e utilized for growth. No growth or only trace of growth on sucrose and raffinose. doubtful .

Color in medium: Melanoid pigments not formed in peptone-yeast- T r a c e s of

One observer r e p o r t s

Utilization of .L-arabinoae is

S t r eptove r t ici 1 lium (Arai, Nakada and Suzuki) BaldaCCi, Farina and Locci. Descriptions: Streptomyces olivoreticuli A r a i e t a l . 1957, 435-441; Streptoverticil l ium olivoreticuli (Arai e t al. ) Baldacci, Farina and Locci 1966, 162. Type strain: LOO (ibid.) = IFM 1018, Ara i , personal communication. ISP description by Group B-1.

Spore chain morphology: Umbellate monoverticillate ( = Streptomyces Section Verticillati, biverticillate, Fig. 144). Mature spore chains generally shor t with 3 to LO spores p e r chain. observed on Pr idham and Gottlieb carbon utilization medium with glucose o r inositol, but good sporulation is usually not seen on yeast- mal t agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 145).

media supporting sporulating a e r i a l growth. tion, the surface is pate yellow on yeast-matt agar , oatmeal a g a r and glycerol-asparagine a g a r and white on sa l t s - s ta rch a g a r .

i s h yellowish brown) on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

Color i n medium: Melanoid pigments a r e formed in peptone-yeast- i r o n a g a r and tryptone-yeast broth, but not in tyrosine agar . No pigment is found in the medium in yeast-malt agar , oatmeal agar , sa l t s - s t a r c h a g a r o r glycerol-asparagine a g a r .

variable. carbon source tested and resu l t s v a r y on duplicate dishes i n the s a m e laboratory. to fluctuation of carbon utilization by ultra-violet i r rad ia ted s t ra ins . This fluctuation appears a l s o to be a charac te r i s t ic of the type strain.

ISP 5105 f r o m T. A r a i as IFM 1018 = LOO.

This morphology may be

Color of colony: Aer ia l m a s s color in the White co lor -ser ies on In the absence of sporula-

Reverse side of colony: No distinctive pigments (colorless to gray-

Carbon utilization: Carbon utilization a p p e a r s to be unusually Reports f r o m three o b s e r v e r s differ on utilization of each

The original description (Arai, e t a t . 1957) d i rec ts attention

Streptomyces olivoviridis Kuchaeva, Krasillnikov, Skryabin and Taptykova. Description: Kuchaeva e t at . 1960, 251 (see a l so Engl. k a n s l . , 222-251). Type strain: INMI 1475 = RIA 661 ( E d . ; type s t ra in ?elected by N.A. Krasil'nikov, personal communication). ISP 5211 f r o m N.A. Krasil 'nikov a s INMI 1475 = RIA 661. by Group B-8.

ISP description

Figure 144. Streptoverticillium olivoreticuli. BIV (umbellate monoverticillate) morphology (X 640) on Pridhamand Gottlieb carbon utilization medium plus inositol, 14 days.

micrograph from 28 day culture on salts-starch agar.

agar, 7 days. '' (b) Flexuousand wavy chains with some tendency toward hooks and spirals; 14 day culture on oatmeal agar. l3

Figure 147. 2. olivoviridis. Spiny spores; electron micrograph from 14 day culture on oatmeal agar. l9

Figure 145. Streptoverticillium olivoreticuli. Smooth spores; electron

Figure 146. S. olivoviridis. 'a) R F spore chains (X 300) on yeast-malt


Spore chain morphology: Section Rectiflexibiles (Fig. 146a). The predominantly flexuous o r wavy spore chains may a l so f o r m occasional hooks and open sp i ra l s ( r a re ) resembling o r ,S morphology (Fig. 146b). Mature spore chains generally long with LO to 50 spores p e r chain. This morphology is seen on yeast-malt agar , oatmeal agar , sa l t s - s ta rch aga r and glycerol-asparagine agar . Spore surface: Spiny (Fig. 147).

yeast-malt agar , oatmeal aga r and sa l t s - s ta rch aga r ; ae r i a l mycelium is poorly developed o r white on glycerol-asparagine agar .

Reverse side of colony: Yellow to yellow brown is usually modified by green (olive brown to yellow green o r grayish green) on yeast-malt agar , oatmeal aga r and sa l t s - s ta rch a g a r ; no distinctive substrate mycelium pigment on glycerol-asparagine agar . pigment i s not a pH indicator.

Color in medium: Melanoid pigments a r e not formed in peptone- yeast-iron aga r o r tyrosine agar . i n tryptone-yeast broth. Yellow o r green pigment is found in the medium in yeast-malt agar , oatmeal aga r and sa l t s - s ta rch agar . is not pH sensitive when tested with 0.05 N NaOH o r HC1.

- D-fructose and rhamnose a r e utilized for growth. t race of growth with sucrose, I-inositol o r raffinose.


Substrate mycelium

Some coloration may appear i n medium

This pigment

Carbon uti1ization:g- Glucose, 4 -arab inose , _D-xylose, _D-maMitot, No growth o r only

Streptomyces phaeochromogenes (Conn) Waksman. Descriptions: Actinomyces pheochromogenus (sic ) Conn 1917a, 3, 16 (also reprinted in 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 chains generally straight chains of LO to 50, o r sometimes more than 50, spores p e r chain. agar , oatmeal agar, sa l t s - s ta rch aga r and glycerol-asparagine agar . Spi ra 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: Aer ia l m a s s color usually i n the Red co lor -ser ies on yeast-malt agar , oatmeal agar , sa l t s - s ta rch aga r and glycerol- asparagine agar . Most representative color tabs f r o m the Red color- s e r i e s were 3ca (pale orange yellow) and 5ca o r 5cb (light yellowish pink to grayish yellowish pink). Aer ia l na ss color may a l so be white o r pate yetlow on oatmeal aga r and pate yellow on sa l t s - s ta rch a g a r and g Lyc e r o 1 -a spa ragine agar .

Reverse side of colony: No distinctive pigments (pale yellow o r grayish yellow on oatmeal aga r and sa l t s - s ta rch agar ; light brown to strong brown on glycerol-asparagine aga r and yeast-malt agar).

i ron agar , tyrosine aga r and tryptone-yeast broth. than t r ace of yellow is found in the medium in yeast-malt agar, oatmeal agar , sa l t s - s ta rch aga r o r glycerol-asparagine agar .

Carbon utilization: 2-Glucose, _L-arabinose, sucrose, _D-xyloae,


This morphology is seen on yeast-malt

Color i n medium: Melanoid pigments a r e formed in peptone-yeast- No pigment other


- i-inositol, _D-mannitol, _D-fructose and rhamnose a r e utilized for growth. Repor ts v a r y on utilization of raffinose.

Streptomyce s phaeopurpureus Shinobu. Description: Shinobu 195 7, 63-67. Type strain: 146 e.). ISP 5125 f r o m R. Shinobu as 146. ISP description by Group B -9 .

Spore chain morphology: Section Rectiflexibiles (Fig. 150). Aer ia l mycelium is usually poorly developed o r absent 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 . Mature spore chains may be observed on other media including Gauze's medium No. 1 where chains of LO to 50 o r m o r e spores may be formed. surface: Smooth (Fig. 151).

co lor -ser ies , but a e r i a l sporulation on yeas t -mal t agar , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r is usually inadequate for accura te spore m a s s color determination.

(light brown, strong brown, o r gray ish reddish 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 yeas t -mal t agar ) . not a pH indicator (or is changed only slightly f r o m brown to yellow brown by 0.05 N HCl).

Color in medium: Melanoid pigments a r e formed in peptone-yeast- i ron agar , tyrosine a g a r and tryptone-yeast broth. Yellow, orange, o r r e d pigment is found in the medium i n yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine agar . slightly pH sensit ive 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, sucrose , _D-xy 1 ose, - i -inositol, _D-mannitol, _D-fructose, rhamnose and raffinose a r e utilized f o r growth.

Spore

Color of colony: Aer ia l m a s s color probably in the Gray o r Red

Reverse side of colony: Yellow to yellow brown is modified by r e d

Substrate mycelium pigment is

This pigment is

Streptomyces phaeoviridis Shinobu. Description: Shinobu 1957, 67- 73. Type strain: 503 k d . ) . ISP 5285 f r o m A. Seino, Kakken Kagaku Go. , Tokyo a s 503. ISP description by Group B -12.

Spore chain morphology: Section Rectiflexibiles (Fig. 152). Mature spore chains generally long with LO to 50 spores p e r chain. morphology is seen on yeas t -mal t agar , oatmeal a g a r and sa l t s - s ta rch agar , but a e r i a l mycelium with rnature spore chains is not formed on glyceroL-asparagine a g a r . Spore surface: Smooth (Fig. 153).

yeast-malt agar , oatmeal a g a r and sa l t s - s ta rch a g a r . be seen on these media and on the poorly sporulated sur face of glycerol- asparagine a g a r . )

Reverse 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 gray ish reddish orange). indicator.

Color in medium: Melanoid pigments a r e formed i n peptone-yeast- i r o n a g a r , tyrosine a g a r and tryptone-yeast broth. Orange o r reddish pigment is found in the medium in yeast-malt agar , oatmeal a g a r and glycerol-asparagine a g a r . tested with 0.05 N NaOH o r HC1.

This

Color of CoLony: Aer ia l m a s s color i n the Gray color-series on (White a l s o may

Substrate mycelium pigment is not a pH

This pigment is not pH sensit ive when

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 21 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


Carbon utilization: _D-Glucose, I=-arabinose, sucrose, _D-xylose, - i -inositol, _D-mannitol, 2 - f ruc tose , rhamnose and raffinose a r e utilized for growth.

S t r e ptomyces platensis Pit tenger and Cottlieb. Descriptions: McGuire, B r i t i s h Pa t . Specification 713, 795, Jan. 31, 19%; Pi t tenger and Cottlieb in Waksman 1961, 258-259. Type strain: M5-5353 (Pittenger, personal communication). as M5-5353.

chains generally LO to 50, o r somet imes m o r e than 50, s p o r e s p e r chain. This morphology is seen on yeast-malt agar , oatmeal a g a r , s a l t s - s ta rch a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 155).

Color of colony: Aer ia l m a s s color in the Gray co lor -ser ies on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and g lycero l -aspar - agine a g a r .

to f o r m black, mois 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.

Reverse side of colony: Grayish yellow on oatmeal a g a r and sa l t s - s t a r c h a g a r ; somet imes modified by red to gray ish yellowish pink o r light reddish brown on yeas t -mal t a g a r and glycerol-asparagine a g a r . Reddish subs t ra te mycelium pigment may be changed to co lor less o r pale yellow by 0.05 N HC1.

yeas t - i ron a g a r , tyrosine a g a r or tryptone-yeast broth. T r a c e s of r e d o r yellow pigments a r e found in the medium in yeas t -mal t a g a r , oatmeat agar , s a l t s - s t a r c h a g a r and glycerol-asparagine a g a r . The reddish pigment is somewhat pH sensitive, changing f r o m grayed r e d to nearly co lor less with 0.05 N HC1.

- D-fructose and raffinose a r e utilized for growth, Utilization of - L-arabinose and _D-xylose i s doubtful. growth with rhamnose.

ISP 5041 f r o m R. C. Pit tenger ISP description by Group B-4.

Spore chain morphology: Section Spi ra les (Fig. 154). Mature spore

Special morphological charac te r i s t ic : Spores frequently coalesce


Carbon utilization: 2 - G lucose, sucrose , i - inos i to 1, _D -mannito 1,

No growth o r only t r a c e of

S t r eptomyce s pseudogri seo lu s Okami and Ume zawa . Description : Okami and Umezawa& Okami, Utahara, Oyagi, Nakamura and Umezawa. 1955, 127-128. Type s t ra in : 534 ( x d . ) . ISP 5026 f r o m Y. Okami a s H-16C = 534 = NIHJ 224.

Spore chain morphology: Section Spi ra les (Fig. 157). Spore chains suggesting morphology may a l s o be common. Mature spore chains moderately short , often with 3 to 10 spores p e r chain, but longer chains a r e formed on suitable media. This morphology is 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 is poorly developed on oatmeal agar and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 158).

Color of colony: Aer ia l m a s s color in the White co lor -s e r i e s on yeas t -mat t a g a r and sa l t s - s ta rch a g a r ; poor sporulation on oatmeal a g a r and glycerol-asparagine a g a r . a s gray ish buff i n the original description by Okami and Umezawa (1955).


The a e r i a l mycelium is descr ibed

Figure 152. 2. phaeoviridis. R F spore chains: (a) (X 485) on oatmeal agar, 14 days. 37 (b) (X 1200) on salts-starch agar.

Figure 153. 2. phaeoviridis. Smooth spores; electron micrograph from 14 day culture on oatmeal agar. ’’

Figure 154. 2. platensis. Spiral spore chains (X 800) on glycerol- asparagine agar, 14 days. 27

Figure 155. S. platensis. Smooth spores; electron micrograph from 7 day cztur-s-starch agar. 31 Tangled masses of spirals a r e associated with hygroscopic droplets.

mas e e s of spores. (a) (X 400) on yeast-malt agar, 18 days (cf.Fig. 155). on oatmeal agar, 14 days.

Figure 156. 2. platensis. Hygroscopic droplets containing coalesced

(b) (X 400)2m oatmeal agar, 18 days. 31 (c) (X 200)


Reverse side of colony: No distinctive pigments (co lor less to gray- i s h yellow o r yellowish brown) on yeas t -mal t agar , oatmeal a g a r , s a l t s - s t a r c h a g a r and glycerol-asparagine agar .

i r o n a g a r , ty ros ine a g a r o r tryptone-yeast broth. t r a c e of yellow, is found i n the medium i n yeas t -mal t a g a r , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

- D-mannitol, ._D-fructose and rhamnose are utilized f o r growth. growth o r only trace of growth with s u c r o s e and r a f f b o s e .

Color in medium: Melanoid pigments are not formed in peptone-yeast. N o pigment, o r only

Carbon utilization: _D-GLucose, &-arabinose, _D-xylose, I-inositot , No

Actinomyces pseudolavendulae Kuchaeva, Krasil 'nikov, Taptikova and Gesheva. Description: Kuchaeva e t a t . 1962, 103-124. Type s t ra in : USSR RIA-744 = IMRU 3440 w. ; Krasil 'nikov, personal cornmunica- tion). description by Group B-LO.

m y c e s pseudolavendulae Kuchaeva e t at. , in a l l de ta i l s with the charac te r iza t ion by study-group A-6 for Strepto- m y c e s lavendulae (Waksman and Cur t i s ) Waksrnan and Henrici, ISP 5069. repor ted i n our previous paper (Shirling and Gottlieb 1968, 138). personal communication f r o m N.A. Krasil 'nikov (April, 1964) gives IMRU 3440 = RLA-744 as the type s t ra in for Actinomyces pseudolavendulae. IMRU 3440 is given a s the type s t ra in for Streptomyces Laven- dulae study of t h i s species. 1962 is regarded h e r e as a n objective synonym for Streptomyces Lavendulae (Waksman and Cur t i s ) Waksman and Henrici 1948; and ISP s t ra in 5213 is equivalent to ISP 5069 Streptomyces Lavendulae 9 . ~

ISP 5213 f r o m N . A . Krasillnikov a s EUA 744 = IMRU 3440. ISP

Remarks : The charac te r iza t ion by ISP study-group B-LO for Actino- ISP s t r a i n 5213 is identical

The charac te r iza t ion for Streptomyces lavendulae, LSP 5069 was A

by Waksman (1961) and was used as the type s t ra in for the ISP Actinomyces pseudolavendutae Kuchaeva et at.

Actinomyces pseudovenexue lae Kuchaeva, Kra sillnikov, Taptikova and Gesheva. Description: Kuchaeva s. 1962, 103-124. Type strain: USSR RIA-742 = IMRU 3774 (ibid. ; Kraeitlnikov, personal communication). ISP 5212 f r o m N.A. Krasil 'nikov as RIA-742 = IMRU 3774. description by Group B-9.

Spore chain morphology: Section Rectiflexibiles. Mature spore 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 seen on yeas t -mal t a g a r , oatmeal agar , sa l t s - s t a r c h a g a r and 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 in the Red c o l o r - s e r i e s (5dc, g ray ish yellowish pink) on yeas t -mal t agar , oa tmeal 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 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 somet imes repor ted for yeas t -mal t a g a r , oat- m e a l a g a r and ea l te -s ta rch a g a r . )

yellow to orange yellow o r Light yellowish brown) on yeas t -mal t agar , oa tmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

i r o n aga-r, tyrosine a g a r and tryptone-yeast broth. yellow pigment may be found in the medium in yeast-malt a g a r , oatmeal a g a r , sa l ta -a ta rch a g a r and glycerol-asparagine agar . This pigment is

ISP

Reverse side of colony: No distinctive pigments (light gray ish

Color in medium: Melanoid pigments a r e formed i n peptone-yeast- T r a c e of grayish

Figure 157. S. pseudogriseolus. Spiral spore chains (X 380) on

Figure 158. 5. seudogriseolus. Smooth spores; electron micrograph

Figure 159. A. pseudovenezuelae. RF spore chains (X 1100) on

Figure 160. A. pseudovenezuelae. Smooth spores; electron micrograph

Waksman's starch agar B, 21 days. '* from 21 day'culture on yeast-malt agar. p 9

glycerol-asparagine agar, 14 days.

from 14 xay culture on glycerol-asparagine agar. '


not pH sensitive when tested with 0.05 N NaOH o r HC1.

- i -inositol, _D-mannitol, _D-fructose, rhamnose and raffinose a r e utilized for growth.

Carbon uti lization: _D-G luco se, &-arabinose, sucrose, _D -xy lo se,

Streptomycee purpeofuscus Yamaguchi and Saburi. Description: Yamaguchi and Saburi 1955, 203-207. Type strain: H-5080 &d2). ISP 5283 f rom T. Yamaguchi as H-5080.

Spore chain morphology: Section Rectif lexibiles (Fig. 161). Mature spore chains generally long, often with more than 50 spores per chain. This morphology i s seen on yeast-malt agar, oatmeal agar, salts-starch agar and glycerol-asparagine agar. Spore surface: Smooth (Fig. 162).

Color of colony: Aerial m a s s color in the Gray color-series on yeast- malt agar, oatmeal agar, salts-starch agar and glycerol-asparagine agar.

Reverse side of colony: Distinctive pigments a r e usually not produced (grayish yellow to olive brown) on yeast-malt agar, oatmeal agar, salts- starch aga r o r glycerol-asparagine agar, but dark brown to reddish brown substrate mycelium pigment may be produced. The original description (Yamaguchi and Saburi, 1955) describes purple to brown reverse mycelium color but notes that this coloration may be lost on repeated transfer.

Color in medium: Melanoid pigments a r e formed in peptone-yeast- iron agar, but only a s t races of brownish black color in tyrosine agar o r tryptone-yeast broth. Traces of red, violet o r yellow pigment may be found in the medium in yeast-malt agar, oatmeal agar and salts- starch agar. N NaOH o r HCl.

utilized for growth. Utilization of _D-fructose i s doubtful. No growth o r only t race of growth with sucrose, i-inositol, _D-maanitcl, rhamnose and raffinose.

ISP description by Group B -12.

This pigment is not pH sensitive when tested with 0.05

Carbon utilization: _D-Glucose, 2-arabinose and _D-xylose a r e

Streptomyces pyridomyceticus Okami and Umezawa. Descriptions and nomenclature: Tentative name with incomplete and tentative des- criptinn in Maeda, Kosaka, Okami and Umezawa 1953, 140; Streptomyces albidofuscus Okami and Umezawa in Okami, Maeda and Umezawa 1954, 55-56; authors' rejection of Streptomyces albidofuscus with assignment of the name Streptomyces pyridomyceticus Okami and Umezawa in Umezawa and Okami 1957, 172. Type strain: 451-A8 (Okami and U m e z a w a h Okami, Maeda and Umezawa 1954, 55). ISP 5024 from Y . 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 yeast- malt agar, oatmeal agar, salts-starch agar, glycerol-asparagine agar and other ISP media a s well as on Hickey and Tresner 's amidex agar. Rare spore chains were short with 3 to LO spores per chain and were generally twisted or curved into hooks resembling incomplete spirals. (Fig. 163), although (Fig. 164). and Umezawa, 1954) a lso note the general absence of good aer ia l mycelium on most media. In the original description loose spirals a r e described a s occurring on the sparse aer ia l mycelium of synthetic media. Spore surface: Smooth (Fig. 165). often interspersed with imperfect spores. This characterist ic i s a lso

spore chains were also sometimes found The original descriptions (Maeda et at., 1963 and Okami

Spores were irregular in size and were

Figure 161. 2. purpeofuscus. spore chains on yeast-malt agar,

Figure 162. 2. purpeofuscus. Smooth spores; electron micrograph

Figure 163. 2. pyridomyceticus. Isolated spore chains forming hooks,

21 days (a, X 300; b, X 1200).23

from 14 day culture on oatmeal agar. 22

incomplete spirals or spirals of only one o r two turns (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 to find on this strain.

Figure 164. - S. pyridomyceticus. spore chains (X 600) on glycerol- asparagine agar, 12 days (see also, Figure 163).20

Figure 165. S. pyridomyceticus. Smooth spores; electron micrograph from 14 zay culture on yeast-malt agar. lo


mentioned in the f i r s t description.

yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and g lycero l -asparagine a g a r a s well a s on other ISP media. a g a r o r oatmeal agar , a e r i a l mycelium is thin and white. description d e s c r i b e s the a e r i a l mycelium as thin and white, somet imes with gray o r brownish gray tinge on Czapek's a g a r .

i s h yellow o r olive brown) on yeas t -mal t 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 .

i ron a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment is found in the medium in yeas t -mal 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 .

L -a ra bin0 s e , sucrose , _D -xy lo s e , L-ino s i t o 1, _D -manni t o 1, ,D - f i'x to s e , rhamnose o r raffinose. carbon sources tested, suggests that P r i d h a m and Gottlieb's basa l medium will not support growth of this strain. (Okami and Umezawa 1954) indicates that _D-glucose, &-arabinose, sucrose , 2 -xylose and _D-fructose can be utilized for growth with Caapek's a g a r as the basa l medium.

Color of colony: Aer ia l mycelium is poorly developed o r absent on

When present on yeas t -mal t The original

Reverse side of colony: No distinctive pigments (colorless o r gray-

Color in medium: Melanoid pigments a r e not formed in peptone-yeast-

Carbon utilization: No growth o r only t r a c e of growth with 2-Gtucose,

Absence of growth with _D-gtucose and a l l other

The original description

-

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.

- R F spore chains (Fig. 166a), and some long chains with te rmina l hooks o r primitive s p i r a l s of wide d iameter (Fig. 166b). 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 seen on yeas t -mat t agar , oatmeal a g a r , sa l t s - s t a r c h a g a r and glycerol-asparagine agar . Spore surface: Smooth (Fig. 167).

5cb, gray ish yellowish pink) on yeast-malt a g a r , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . sporulation may a l s o be seen.

Reverse side of colony: No distinctive pigments on oatmeal a g a r , but y e l l w t o yellow browp subs t ra te mycelium color may be modified i n some a r e a s by blue pigment (dark olive green, gray ish green o r grayish blue) on yeas t -mal t a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . be intensified f r o m grayish green to grayish blue o r violet by 0.05 N NaOH o r f r o m gray green to reddish gray by 0.05 N HC1.

i ron agar , tyrosine a g a r and tryptone-yeast broth. than a t r a c e of yellow is found in the medium i n yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine agar .

for growth. Q-mannitot, _D-fructose, rhamnose and raffinose.

ISP description by Group B-4. Spore chain morphology: Section Retinaculiaperti including many

Mature spore chains

Color of colony: A e r i a l m a s s color in the Red co lor -ser ies (5ec and

A r e a s of g r a y o r white

The blue pigment is not always present. When present , i t can

Color in medium: Melanoid pigments are produced i n peptone-yeast- No pigment other

Carbon utilization: _D-Glucose, 5 -arab inose and sucrose a r e utilized Only a t r a c e of growth is found on _D-xylose, -inositol,

Figure 166. 2. racemochromogenus. morphology including both straight spore chains and terminal spirals: (a) R.F chains (X 400) on yeast-malt agar, 17 days. 31 (b) T y p i c a l 2 terminal hooks and spirals (X 200) on oatmeal agar, 14 days."

Figure 167. 2. racemochromogenus. Smooth spores; electron micrograph from 14 day culture on oatmeal agar. 40

Figure 168. S. rimosus. Spiral spore chains (X 1100) from,14 day c u l t u r e s ; n : p a s t - m a l t agar, (b) oatmeal agar, (c) glycerol- asparagine agar.

Figure 169. 3. rimosus. Smooth spores; electron micrograph from 14 day culture on yeast-malt agar.'


Streptomyces r imosus (Sobin, Finlay and Kane) Waksman. Descrip- tions: Sobin, Finlay and Kane, U.S. Pa t . 2,516,080 July 18, 1950. Waksman & W a k s k n and Lechevalier 1953, 47. Type -strain: Pf izer F.D. 10326 = NRRL 2234, IMRU 3558, ATCC 10970 (Routien, personal communication). ISP 5260 f r o m J.B. Routien, Chas. Pf izer and Co. , Inc. a s FD 10326.

imperfec t sp i ra l s , hooks, and Loops suggesting present . o r often m o r e than LO, s p o r e s p e r chain. yeas t -mat t agar , oatmeal agar , sa l t s - s ta rch a g a r , glycerol-asparagine a g a r and on carbon utilization medium plus r)-glucose. Smooth (Fig. 169).

on yeast-malt a g a r and in the Yellow o r White co lor -ser ies on oa tmeal 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 are 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 co lor - wheel.

Reverse s ide of colony: Grayish yellow on oatmeal a g a r and sa l t s - s t a r c h a g a r ; g ray ish yellow modified slightly by r e d (modera te to dark orange-yellow) on yeas t -mal t a g a r and glycerol-asparagine agar . s t r a t e mycelium pigment is not a pH indicator.

yeast-iron agar , tyrosine a g a r o r tryptone-yeast broth. yellow pigment is found in the medium in yeas t -mal t 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 . pH sensit ive when tested with 0.05 N NaOH o r H C l .

- D-fructose and raffiaoee a r e utilized for growth. xylose is doubtful. and raffinose.

ISP description by Group B-9. Spore chain morphology: Section Spi ra les (Fig. 168), but many

morphology a r e a l so Mature spore chains shor t to moderately long with 3 to LO,

This morphology is seen on

Spore surface:

Color of colony: Aer ia l m a s s color in the Red or White c o l o r - s e r i e s

Sub-

Color in medium: Melanoid pigments a r e not formed i n peptone- T r a c e of

This pigment is not

Carbon utilization: _D-Gtucose, &-arabinose, i- inositol , 2- mannitol, Utilization of _D-

No growth o r only t r a c e of growth with sucrose

Streptomyces rochei B e r g e r , Jampolsky and Goldberg. Description: Name only, without description is found in B e r g e r , Jampolsky and Coldberg 1949, 476-478. 1953, 40-41 and by Preobrazhenskaya Et 51. & Gauze ct a_l. 1957, 133 (Engl. t r a n s l . , 106). Type strain: X 15 (Berger , personal communication). ISP 5231 f r o m J. Berger , Hoffman-LaRoche, Inc. as X 15. ISP description by Group B-8.

Spore chain morphology: Section Spi ra les (Fig. 170a). Spi ra l s a r e usually open, somet imes a lmost flexuous (Fig. 170 b, c). chains moderately tong with LO to 50, o r often m o r e than 50 spores 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 . Spore surface: Smooth to warty (Fig. 171).

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- mat t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

Reverse side of colony: No distinctive pigments (grayish yellow to yellowish brown) on yeas t -mat t 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 .

Description is given in Waksman and Lechevalier

Mature spore

Figure 170. 2. rochei. (a) Spiral spore chains (X 300) on glycerol- asparagine agar, 7 d a y s T b , c) Open spirals to flexuous chains (X 400) on oatmeal agar, 14 days. 41

from 14 day culture on oatmeal agar. l 9

Figure 171. 5. rochei. Smooth to warty spores; electron micrograph


Color in medium: Melanoid pigments a r e not formed in peptone-yeast- i ron a g a r , tyrosine a g a r o r tryptone-yeast broth. N o pigment i s found in the medium in yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

Carbon utilization: _D-GLucose, _L-arabino se, 2-xylose , i -inositol, - D-mannitol, _D-fructose and rhamnose a r e utilized for growth. growth or only t r a c e of growth on sucrose and raffinose.

No

Streptomyces roseo-luteus Besse t t . Description: Besse l l , U.S. Patent 3,013, 947 Dec. 19, 1961). Type s t ra in : NRRL 2776 = NCIB 8984 (ifi.). ISP description by Group B-6.

many turns occur at ends of long spore chains (Fig. 172). t imes sugges tRA morphology. with LO to 50, o r often m o r e than 50, spores p e r chain. This morphology i s seen on yeas t -mal t 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. 173).

yeast-malt agar , oatmeal a g a r , sa l t s - s ta rch a g a r and g lycero l -aspar - agine a g a r . with color tab 5fe (light gray ish reddish brown) as the n e a r e s t matching color f r o m the Gray color wheel. c o l o r - s e r i e s were 5ge and 4ge (light grayish reddish brown).

oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r ; subs t ra te mycelium pigment is not a pH indicator.

i ron agar , tyrosine a g a r o r tryptone-yeast broth. Yellow pigment is found in the medium in yeast-matt 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 not pH sensit ive 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 for growth. with _L-arabinose, sucrose , 1 -inositol and rhamnose.

ISP 5240 f r o m C. W. Hesselt ine (NRRL) as NRRL 2776.

Spore chain morphology: Section Sgi ra tes . Tight s p i r a l s of one to These some-

Mature spore chains generally long

Color of colony: Aer ia l m a s s color in the Red c o l o r - s e r i e s on

One observer placed this s t ra in in the Gray c o l o r - s e r i e s

Neares t matching tabs i n the Red

Reverse side of colony: Yellow to orange yellow on yeas t -mat t a g a r ,


No growth o r only t r a c e of growth

Streptomyces roseosporus Fa lczo de Morais and Dalid Maia. Description: Streptomyces venezuelae var . roseospor i Fa lczo de Morais, Dalid h4aia and Genn 1958, 102-103. Streptomyces roseosporus Morais and D. Maia (,,c, nom nov 1961, 43. Type strain: 4192 (Falcxo de Morais and Dalid Maia 1961, 43). 5122 f r o m J.O. FalcZo de Morais as 4192.

Spore chain morphology: Section Rectiflexibiles. Mature spore chains generally long, often with m o r e than 50 s p o r e s p e r chain (Fig. 174). s t a r c h a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 175).

mal t agar , oatmeal a g a r and s a l t s - s t a r c h a g a r . co lor -ser ies on glycerol-asparagine a g a r . )

Reverse side of colony: No distinctive pigments (grayish yellow to yellowish brown) 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 .

Change in name (rank):

ISP ISP description by Group B - 3 .

This morphology is seen on yeas t -mal t agar , oatmeal a g a r , sa l t s -

Color of colony: Aerial m a s s color in the Red c o l o r - s e r i e s on yeas t - (White, Yellow o r Red

Figure 172. S. roseo-luteus. Spiral spore chains (X 500) on salts-starch

Figure 173. 2. roseo-luteus. Smooth spores (Au/Pd shadowed); electron

Figure 174. S . roseosporus. RF spore chains.

Figure 175. 2. roseosporus. Smooth spores; electron micrograph from

agar, 3 X a y s 7 -

micrograph from 21 day culture on salts-starch agar. 44

agar, 14 days. lo (b) (X 600) on glycerol-asparagine agar, 21 days. 2o

14 day culture on salts-starch agar.

(a) (X 300) on oatmeal

3 7 2 I N T E R N A T I O N A L J O U R N A L

Color in medium: Melanoid pigments a r e not formed in peptone- yeast-iron a g a r , tyrosine a g a r o r tryptone-yeast broth. No pigment is found in the medium in yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine agar .

Carbon utilization: _DGlucose, ,L -arabinose, 2 -xylose and rhamnose a r e utilized f o r growth.

o r only t r a c e of growth with sucrose , t -inositol , D-mannitol aad raffinose.

Utilization of _D-fructose is doubtful. No growth

-

Actinomyces roseoviridis Preobrazhenskaya. Description: P r e o - brazhenskaya @ Gauze ct 51. 1957, 57. Type strain: 3617 (Preobrazhen- skaya, personal communication; G. F. Gauze, cited by Gottlieb 1968, 20). Streptomyces roseoviridis (Preobrazhenskaya) Pr idham 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 spore chains generally LO to 50 s p o r e s p e r chain. seen on yeast-malt agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine a g a r . Spore surface: Smooth (Fig. 177).

on yeas t -mal t a g a r , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- a sparagine a g a r .

oatmeal agar , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

i ron a g a r , tyrosine a g a r and tryptone-yeast broth. P igments other than melanoids a r e not formed in yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r .

for growth. - D-mannitol, rhamnose and raffinose. Utilization of _D-fructose is doubtful.

This morphology is

Color of colony: A e r i a l m a s s color in the Red o r Yellow color -ser ies

Reverse side of colony: No distinctive pigment on yeas t -mal t a g a r ,

Color in medium: Melanoid pigments a r e formed i n peptone-yeast-

Carbon utilization: _D-GLucose, _L-arabinose and _D-xylose a r e utilized No growth o r only t r a c e of growth on sucrose , -inositol,

Actinomyces rubiginosohelvolus Kudrina. Description: Kudrina in Gauze et a l . 1957, 78 and 89. Type s t ra in : 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 as 10/53.

Spore chain morphology: Section Rectiflexibiles (Fig. 178). Aer ia l mycelium is somet imes poorly developed on ISP media. Mature spore chains, when formed, contain LO to 50 o r m o r e s p o r e s p e r chain on yeas t -mal t a g a r , oatmeal agar , sa l t s - s ta rch a g a r and g lycero l -aspar - agine a g a r . Spore surface: Smooth (Fig. 179).

Color of colony: Aer ia l m a s s color in the Yellow c o l o r - s e r i e s on yeas t -mal t a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . Aer ia l mycelium is poorly developed on oatmeal a g a r . One observer a l s o recorded tabs 2ca(pale yellow) and 3ca (pale orange yellow) f r o m the Red co lor -ser ies as a second color on yeas t -mal t a g a r and glycerol- asparagine a g a r .

on yeas t -mal t agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol- asparagine a g a r . to r changing f r o m yellow to pink with addition 0.05 N NaOH.

ISP description by Group A-12.

Reverse side of colony: Colorless to pale yellow o r grayed yellow

However, yellowish r e v e r s e pigment is a pH indica-

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

Figure 178. _A. rubiginosohelvolus. RF spore chains (X 400) on yeast-

Figure 179. _A. rubiginosohelvolus. Smooth spores; electron micrograph

21 day c & u r e on yeast-malt agar. 23

malt agar, 21 days.z3

from 21 day culture on yeast-malt agar. 23


Color i n medium: Melanoid pigments not formed in peptone-yeast- i ron agar , tyrosine a g a r o r tryptone-yeast broth. No pigments o r only t r a c e of yellow pigment found in yeast-malt agar , oatmeal a g a r , sa l t s - s t a r c h a g a r o r glycerol-asparagine a g a r .

- D-fructose and rhamnose a r e utilized for growth. t r a c e of growth on sucrose, i - inos i to l and raffinose.

Carbon uti t i zation : _D -G luc o s e, 4 -a rabino s e , ,D -xy Lo s e , _D - mannito 1, No growth o r only

Actinomyces rubiginosus Preobrazhenskaya, B linov and Ryabova. Description: Preobrazhenskaya et al . in Gauze e t a t . 1957, 132 and 134. Type s t ra in : 11852 (Gauze & Cotttieb, 1968). Streptomyces rubiginosus (Preobrazhenskaya, Blinov and Ryabova) P r i d h a m e t a l . 1958, 70. ISP 5177 f r o m T.P. Preobrazhenskaya as 11852. ISP description by Group A-12.

on s a l t s - s t a r c h a g a r . Aer ia l mycelium and spore chains may be poorly developed and atypical on yeas t -mal t a g a r , oatmeal a g a r and glycerot- asparagine a g a r (Fig. 180). spores p e r chain. Spore surface: Spiny (Fig. 181).

yeas t -mat t agar , oatmeal a g a r , sa l t s - s ta rch a g a r and g lycero l -aspar - agine a g a r .

grayed orange brown o r yellowish gray) on yeast-malt agar , oatmeal agar , s a l t s - s t a r c h agar and glycerol-asparagine agar .

i ron agar , tyrosine a g a r o r tryptone-yeast broth. in medium in yeast-malt a g a r , sa l t s - s ta rch a g a r o r glycerol-asparagine a g a r . this pigment is not pH sensitive.

- D-mannitol, _D-fructose and rhamnose a r e utilized f o r growth. on sucrose i s l e s s abundant than on other carbon sources . o r only t r a c e of growth on raffinose.

Spore chain morphology: Section Spirales. Spi ra l s a r e bes t developed

Mature spore chains generally LO to 50

Color of colony: Aer ia l mass color in the Gray co lor -ser ies on

Reverse side of colony: No distinctive pigment (yellowish brown,

Color in medium: Melanoid pigments not formed in peptone yeas t - No pigment is found

There may be a t r a c e of r e d soluble pigment i n oatmeal a g a r ;

Carbon utilization: _D-Glucose, sucrose , _D-xylose, i -inositol, Growth

No growth

Streptomyces scabies (Thaxter) Waksman and Henrici . Descriptions: Oospora scabies Thaxter 1892, 159-160. Actinomyces scabies (Thaxter) Giissow 1914, 433; Millard and B u r r 1926, 628- 629. Streptomyces scabies (Thaxter) Waksman and Henrici 1948, 957. Type strain: IMRU 3018 (Waksman 1961, 274-275). 3018. ISP description by Group B -4.

spore chains moderately shor t with 3 to LO, o r somet imes m o r e than 10, spores p e r chain. oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . Spore surface: Smooth (Fig. 183).

yeas t -mal t a g a r , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

by green to light gray ish olive, olive, o r olive brown on yeas t -mat t agar , oatmeal agar , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . Substrate mycelium pigment is not a pH indicator.

ISP 5078 f r o m S.A. Waksman a s IMRU


This morphology is seen on yeas t -mat t agar ,

Color of colony: Aer ia l m a s s color in the Gray c o l o r - s e r i e s on

Reverse side of colony: Yellow to yellow brown is usually modified

Figure 180. A. rubiginoeus. Spiral spore chains; imperfect spirals (X 400) on yeast-malt agar, 21 days."

Figure 181. _A. rubiginosus. Spiny spores; electron micrograph from 21 day culture on oatmeal agar. ''

Figure 182. S. scabies. RF epore chains: (a) (X 400) on yeast-malt agar, 17-days. 31

Figure 183. 5. scabies. Smooth spores; electron micrograph from 14 day culture on oatmeal agar. ''

Figure 184. S. sindenensis. - RF spore chains (X 200) on oatmeal agar, 14 days.m

Figure 185. 2. sindenensis. Long chains of smooth spores; electron micrograph (X 2700) from 16 day culture on yeast-malt agar. ''

Figure 186. 2. sindenensis. Smooth spores (X 20,000); electron micrograph from 14 day culture on yeast-malt agar.

-

(b)(X 800) on salts-starch agar, 14 days. ''


Color i n medium: Melanoid pigments a r e not formed in peptone- yeast-iron agar , tyrosine a g a r o r tryptone-yeast broth. Yellow o r greenish yellow pigment may be found in the medium in yeas t -mal t a g a r and oatmeal agar , but not in sa l t s - s ta rch a g a r o r g lycero l -aspar - agine a g a r ; t h i s pigment is not pH sensit ive when tested with 0.05 N NaOH o r H C l .

- D-fruc tose and rhamnose a r e utilized for growth. t r a c e of growth with sucrose ,

Carbon utilization: 2-G lucose, &-arabinose, _D-xylo se, _D -mannitol , No growth or only

i -inositol and raffinose. - Streptomyces sindenensis Nakazawa and Fujii. Descriptions:

Nakazawa and Fujii 1957, 109-110; Nakazawa 1964, 69-70. Name only, without description, in Nakazawa, Fujii , Inoue, Hitomi, Miyake and Kaneko 1954, 168. Type s t ra in : 1071 ( i . . ) . ISP 5255 f r o m K . Nakazawa a s 1071 = IF0 3399.

spore chains short , often with only 3 to LO s p o r e s p e r chain (longer chains may be found on impress ion mounts o r electron micrographs . Fig. 185). This morphology i s seen on yeas t -mal t 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. 186).

sporutating a e r i a l mycelium is poorly developed on ISP media. When sporulation occurs, i t is usually in the White o r Red (5cb, grayish yellowish pink) co lor -ser ies on yeas t -mal t a g a r and in 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 in the White co lor -ser ies on sa l t s - s ta rch a g a r and glycerol- asparagine a g a r .

Reverse side of colony: No distinctive pigments (colorless o r pale grayish yellow) 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 g tycerol -asparagine a g a r .

i ron 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 agar , sa l t s - s ta rch a g a r o r g Lyce r o 1 -a spa ragine a g a r .

and 2 - f r u c t o s e a r e utilized for growth. growth with sucrose , i -inositol, rhamnose and raffinose.

ISP description by Group B -11. Spore chain morphology: Section Rectiflexibiles (Fig. 184). Mature

Color of colony: Aer ia l m a s s color i s difficult to determine because


Carbon utilization: _D-GLucose, _L-arabinose, _D-xylose, _D-mannitol No growth o r only t r a c e of

Streptomyces somaliensis (Brumpt) Waksman and Henrici . Descrip- tion: Indiella somaliensis Brumpt 1906, 555 and 561-562. See also: Actinomyces somaliensis Er ikson 1935, 17-18; Streptomyces somalien- - sis (Brumpt) Waksman and Henrici 1948, 965. (Erikson 1935, 17, neotype). ISP 5267 f r o m S . T. Cowan, NCTC a s NCTC 3236.

most common (Fig. 187). somet imes m o r e than 50. s p o r e s p e r chain. on yeas t -mat t agar , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol- asparagine a g a r . Spore surface: Smooth (Fig. 188).

Color of colony: Aer ia l m a s s color in the Yellow c o l o r - s e r i e s (2db o r Zba, pale yellow) 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 glycerol-asparagine a g a r . (One observer selected color tab

Type s t ra in : 3236

ISP description by Group B-11. Spore chain morphology: Section Rectiflexibiles; flexuous chains a r e

Mature spore chains generally LO to 50, o r This morphology i s seen

Figure 187. S. somaliensis. &F spore chains on oatmeal agar, 14 days. 36

Figure 188. S. somaliensis. Smooth spores; electron micrograph from

Figure 189. S. s heroides. Spiral spore chains (X ca 900) on salts-

Figure 190. 5. spheroides. Smooth spores; electron micrograph from

- 14 day culture on yeast-malt agar.

starch agar, '14 days. * - 14 day culture on salts-starch agar.


3ca, pale orange yellow, f rom the Red color wheel a s the most representative color.) Aerial mycelium is often poorly developed on much of the culture surface on ISP media. color-series was observed on Czapekls agar .

grayish greenish yellow and light olive brown) on yeast-malt agar, salts- s tarch agar and glycerol-asparagine agar.

yeast-iron agar, tyrosine agar o r tryptone-yeast broth. is found in the medium in yeast-matt agar , oatmeal agar , salts-starch agar and glycerol-asparagine agar; this pigment is not pH sensitive when tested with 0.05 N NaOH or HC1.

- D-mannitol and E-frucGse a r e utilized for growth. t race of growth with i-inositol, rhamnose o r raffinose.

Good sporulation in the Yellow

Reverse side of colony: No distinctive pigments (grayish yellow to

Color in medium: Melanoid pigments a r e not formed in peptone- Yellow pigment

Carbon utilization: D-Glucose, _L-arabinose, sucrose, _D-xylose, No growth o r only

- Streptomyces spheroides Wa llick, Harris, Reagen, Ruger and

Woodruff. Description: Wallick et at . 1956, 909-917. Type strain: MA 319 (Woodruff, personal communication 1964). ISP 5292 from H. B. Woodruff, Merck, Sharp and Dohme a s MA 319 ( =NRRL 2449). ISP description by Group B-12.

Spore chain morphology: Section Spirales (Fig. 189). Mature spore chains generally long, often with more than 50 spores per chain, morphology is seen on yeast-malt agar, oatmeal agar , sal ts-s tarch agar and glycerol-asparagine agar. Spore surface: Smooth (Fig. 190).

yeast-malt agar , oatmeal agar , salts-starch aga r and glycerol-asparagine agar .

yellow o r light yellowish brown) on yeast-malt agar , oatmeal agar, salts-starch agar and glycerol-asparagine agar.

yeast-iron agar, tyrosine agar or tryptone-yeast broth. No pigment is found in the medium in yeast-malt agar, oatmeal agar, salts-starch agar o r glycerol-asparagine agar.

- D-mannitol, 2-fructose and rhamnose a r e utilized for growth. Only a t race of growth is found on 'i-inositol and raffinose. A t race of growth also occurs on the carbon-free basal medium.

This

Color of colony: Aerial mass color in the Yellow color-series on

Reverse side of colony: No distinctive pigments (colorless to pale

Color in medium: Melanoid pigments a r e not forlned in peptone-

Carbon utilization: _D-Glucose, _L -arabinose, sucrose, _D-xylose,

Actinomyce s streptomycini Krasil'nikov. Descriptions : Actinomyce s globisporus streptomycini Krasilanikov 1949, 100; Actinomyces strepto- myciniKrasil'nikov 1955, 12-18; see also Nikitina et at. 1960, 85-103 (Engl. transl . 80-98). Type strain: RIA 333 = INh4I 1448 (Krasil'nikov, personal communication, 1964). RLA 333 = INMX 1448 ( =ATCC 15886). for Streptomyces gr iseus (Krainsky) Waksman and Henrici, ISP 5236 = IMRU 3463 1.v. ISP description by Group B -3.

a r e most common (Fig. 191). often with more than 50 spores per chain. This morphology i s seen on yeast-malt agar, oatmeal agar, salts-starch agar and glycerol-asparagine agar . Spore surface: Smooth (Fig. 192).

ISP 5200 from N.A. Krasilanikov a s Note: This is a subjective synonym

Spore c h i p morphology: Section Rectiflexibiles, flexuous chains Mature spo;e chains generally long,

Figure 191. A. streptomycini. RF spore chains (X 300) on yeast-malt

Figure 192. A. streptomycini. Smooth spores; electron micrograph

Figure 193. 2. tanashiensis. RF spore chains (X 312) on glycerol-

Figure 194. S. tanashiensis. Smooth spores; electron micrograph from

agar, 15days. "

from 14 day culture on yeast-malt agar. I t

asparagine agar, 14 days?

10 day culture on oatmeal agar.


Color of colony: A e r i a l mass color in the Yellow c o l o r - s e r i e s on yeas t -mal 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 ( somet imes in the White c o l o r - s e r i e s on glycerol- a sparagine agar ) .

yellow o r light olive brown) on yeas t -mal t 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 .

i r o n a g a r , ty ros ine a g a r o r tryptone-yeast broth. No pigment is found in the medium i n yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r o r g lycerol-asparagine a g a r .

Carbon uti Liza tion : _O -G lu c o s e, _D -xy lose, _D - ma nnit o 1 and _D - f r u c to s e and utilized f o r growth. arabinose, sucrose , , i-inositol, rhamnose and raffinose.

Reverse side of colony: No distinctive pigments (colorless, grayish


No growth o r only t r a c e of growth with 4- -

Streptomyces tanashiensis Hata, Ohki and Higuchi. Descriptions: Hata, Ohki and Higuchi 1952a, 529-534; 1%2b, 67-77 (448-457); 1952c, 188-189. Type s t ra in : 144 (ibid.). ISP 5195 f r o m Y. Yagisawa f r o m T. Hata,

spore chains generally LO to.50, o r somet imes m o r e than 50, s p o r e s p e r chain. sa l t s - s ta rch a g a r and glycerol-asparagine a g a r . Smooth (Fig. 194).

yeas t -mal t agar , oa tmeal a g a r , sa l t s - s ta rch a g a r and g lycero l -asparagine a g a r .

Reverse side of colony: No distinctive pigments (pale yellow to light olive brown) on yeas t -mal 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 formed in peptone-yeast- i r o n a g a r and tryptone-yeast broth, but not on tyrosine a g a r . No pigment is found in the medium i n yeas t -mal t agar , oatmeal a g a r , sa l t s - s t a r c h a g a r o r glycerol-asparagine a g a r .

utilized for growth. Repor ts v a r y on utilization of _D-fructose. Only a t r a c e of growth is found with sucrose , rhamnose and raffinose.

Kitasato Inst . , as 144. ISP description by Group B-5. Spore chain morphology: Section Rectiflexibiles (Fig. 193). Mature

This morphology is seen on yeas t -mal t a g a r , oatmeal a g a r , Spore surface:

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

Carbon utilization: _D-Glucose, _L-arabinose and _D-xylose a r e

i-inositol, _D-rnannitol, -

Actinomyces violascens Preobrazhenskaya and Sveshnikova. Des- cription: Preobrazhenskaya a n d Sveshnikova & Gauze et at. 1957, 32 and 41. Type strain: 3959/54 (Ibid.; see a l s o Gauze&Gottl ieb 1968). S t rep tomyces violascens (Preobrazhenskaya and Sveshnikova) P r i d h a m %tat. 1958, 68. ISP 5183 f r o m T.P . Preobrazhenskaya a s 3959/54.

LO t o 50 o r m o r e s p o r e s p e r chain. 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 of colony: Aer ia l m a s s color in the Violet c o l o r - s e r i e s on sa l t s - s ta rch a g a r and glycerol-asparagine a g a r ; Violet o r White color- series on oatmeal a g a r ; Violet, White o r Red c o l o r - s e r i e s on yeas t - malt agar .

ISP description by Group A-13. Spore morphology: Section Spi ra les (Fig. 195). Mature spore chains

This morphology is seen on yeas t -

Spore sqface: Spiny (Fig. 196).

Figure 195. A. violascens. Spiral spore chains (X 500) on yeast-malt

Figure 196. 4. violascens. Spiny spores; electron micrograph from agar, 24days. l5

24 day culture on yeast-malt agar. 24


Reverse side of colony: No distinctive pigment (grayed yellow to yellow-brown) on yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

i ron agar , tyrosine a g a r and tryptone-yeast broth. P igments other thac melanoids not formed (or only t r a c e s of yellow) in yeas t -mal t a g a r , oatmeal a g a r , sa l t s - s ta rch a g a r and glycerol-asparagine a g a r .

and raffinose a r e utilized for growth. on _D-mannitol and rhamnose. doubtful.

Color in medium: Melanoid pigments a r e formed in peptone-yeast-

Carbon uti lization: _D -G luco se, 4 -a rabino se, _D -xy lo s e, ,D -f ructo s e No growth o r only t r a c e of growth

Utilization of sucrose and--inositoL is

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Documents

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