International Journal of Systematic Bacteriology (1 999), 49, 179-1 84 Printed in Great Britain

Polyamine profiles within genera of the class NOTE with u-diaminopimelic acid in the peptidoglycan

Hans-Jurgen Busse't and Peter Schumann2

Author for correspondence: Hans-Jurgen Busse. Tel: +43 1 25077 2128. Fax: +43 1 25077 2190. e-mail : Hans-Juergen. Busse @vu-wien.ac.at

1 Institute of Microbiology Polyamine patterns of coryne- and nocardioform representatives of the class and Genetics, University of Actinobacteria with u-diaminopimelic acid in the peptidoglycan, comprising Vienna, A-1030 Vienna, Austria strains of the genera A eromicrobium, Nocardioides, In trasporangium, Terrabacter, Terracoccus, Propioniferax, Friedmanniella, Microlunatus, * DSMZ-German Collection of Microorganisms and Luteococcus and Sporichthya, were analysed. The different polyamine patterns Cell Cultures GmbH, were in good agreement with the phylogenetic heterogeneity within this D-07745 Jena, Germany group of actinomycetes. Strains of the closely related genera Nocardioides and Aeromicrobium were characterized by the presence of cadaverine. The second cluster, consisting of the type strains of the Friedmanniella antarctica, , Microlunatus phosphovorus and Luteococcus japonicus, displayed as a common feature the presence of the two predominant compounds spermidine and spermine. The presence of putrescine was common to the type strains of the species Intrasporangium calvum, Terrabacter tumescens and Terracoccus luteus. Sporichthyapolymotpha, which is a representative of a separate line of descent, displayed spermidine as the predominant polyamine. These data indicate that polyamine patterns are suitable for the classification of actinomycetes with LL-diaminopimelic acid in the peptidoglycan.

Keywords : polyamine patterns, chemotaxonomic classification, LL-diaminopimelic- acid-containing Actinobacteria

Coryneform and nocardioform actinomycetes with LL- distinctly separated from the three clusters of diaminopimelic acid (LL-A,~~)as diagnostic diamino coryneform and nocardioform actinomycetes, which acid of their peptidoglycan are a biotechnologically comprise the following genera : (i) Nocardioides and interesting group of metabolically versatile . Aeromicrobium (Collins et al., 1994; Collins & Representatives of this group have, for example, the Stackebrandt, 1989; Miller et al., 1991 ; O'Donnell et capacity to accumulate polyphosphate (Nakamura et al., 1982; Tamura & Yokota, 1994) (Nocardioidaceae); al., 1995), to produce the antibiotic erythromycin (ii) Terrabacter (Collins et al., 1989), Terracoccus (Miller et al., 1991) and to transform steroids (Prauser et al., 1997) and Intrasporangium (Hayakawa & Kurokawa, 1963; Nobile et al., 1955). (Kalakoutskii, 1989) (Intrasporangiaceae); (iii) Organisms with LL-A,~~occupy different phylogen- Propioniferax (Yokota et al., 1994), Luteococcus etic lineages which represent five families within the (Tamura et al., 1994), Microlunatus (Nakamura et al., class Actinobacteria, as revealed by 16s rRNA se- 1995) and Friedrnanniella (Schumann et al., 1997) quence analyses (Prauser et al., 1997; Stackebrandt et (). These genera are distinguished al., 1997). The sporoactinomycetes Streptomyces from each other by their chemotaxonomic charac- (Streptomycetaceae) (Stackebrandt et al., 199 1) and teristics. The genera Nocardioides, Aeromicrobium, Sporichthya (Sporichthyaceae)(Rainey et al., 1993) are Luteococcus and Propioniferax share the peptido- glycan type A3y (Schleifer & Kandler, 1972), with a *Present address: lnstitut fur Bakteriologie, Mykologie und Hygiene, single glycine residue as interpeptide bridge. The Veterinarmedizinische Universitat Wien, Veterinarplatz 1, A-I 210 Wien, genera Terrabacter, Terracoccus and Intrasporangium Austria. are characterized by the cross-linkage of the A3y-type Abbreviation : LL-A,pm, LL-diaminopimelicacid. peptidoglycan via a triglycine-interpeptide bridge and

00717 0 1999 IUMS 179 H.-J. Busse and P. Schumann by a glycine residue bound to the a-carboxyl group of Corynebacterium (Altenburger et al., 1997). When the D-glutamic acid at position 2 of the peptide subunit. Nocardiodes albus strains IMET 7807T and IMET The genera Microlunatus and Friedmanniella are dis- 78 15 were grown on a mineral medium (Lindenbein, tinguished by the rare peptidoglycan type A3y' 1952), cadaverine was detected as the predominant (Schleifer & Seidl, 1985), with glycine at position 1 of compound. The high concentration of putrescine in the peptide subunit. The genera Nocardioides, the majority of strains of the genus Nocardioides when Terrabacter and Terracoccus have MK-8(H4) as their grown in R medium, in addition to cadaverine, appears major isoprenoid quinone, the genus Intrasporangium to be useful for differentiation from the genus shows MK-8, whereas the menaquinone MK-9(H4) is Aeromicrobium. The two species Aeromicrobium common to all other coryne- and nocardioform actino- fastidiosum and Aerornicrobium erythreum are phylo- mycetes with LL-A,pm. Fatty acid- and polar lipid genetically associated with the genus Nocardioides patterns are useful tools for the further differentiation (Tamura & Yokota, 1994) and are included in the of this group of organisms (Schumann et al., 1997). family Nocardioidaceae (Stackebrandt et al., 1997). Recently, the suitability of polyamine patterns for The polyamine patterns of these two species were classification of coryneform actinobacteria has been similar to members of the genus Nocardioides in demonstrated (Altenburger et al., 1997 ; Hamana, containing significant concentrations of cadaverine, 1995, 1996). However, these studies did not include which distinguished A erom icrob iumfas t idiosum I M ET representatives of the class Actinobacteria with LL- 7849T and Aeromicrobium erythreum NRRL B-338 lT A,pm or the data are difficult to interpret due to from the other LL-A,pm-containing actinobacteria. nonstandardized growth conditions. Thus, we However, the two species did not display identical analysed the polyamine composition of coryne- and polyamine patterns. Aerornicrobium fastidiosum IMET nocardioform actinomycetes with LL-A,pm to estimate 7849T contained a rather low concentration of poly- the value of this approach for classification of members amines and spermine was found to be the predominant of this phylogenetically heterogeneous group compound, whereas Aeromicrobium erythreum NRRL (Schumann et al., 1997). B-3381T was characterized by the second major com- pound, spermidine. Strains analysed are listed in Table 1. Biomass for polyamine analyses was obtained by submerged cul- The second polyamine similarity cluster consisted of tivation of the strains in R medium (Yamada & the phylogenetically closely related species Propioni- Komagata, 1972) or polyamine-free glucose/ ferax innocua, Microlunatus phosphovorus and Luteo- asparagine medium (MM) (Lindenbein, 1952) at 28 "C coccus japonicus, which are included in the family using a horizontal shaker run at 220 r.p.m. Cells were Propionibacteriaceae, and Friedmanniella antarctica harvested in the exponential phase of growth when (Schumann et al., 1997). Due to its phylogenetic 70% of the maximal optical density at 578 nm was position and similar chemotaxonomic characteristics reached. Because of the tendency to aggregation of Friedmanniella antarctica also belongs to the family Luteococcus japonicus IF0 12422T and Propioniferax Propionibacteriaceae (Schumann et al., 1997 ; innocua NCTC 11082T, their cultivation times were Stackebrandt et al., 1997). The type strains of these estimated. Sporichthya polymorpha IMET 7840T and four species, NCTC 11082T,JCM 9379T, IF0 12422T Friedmanniella antarctica AA-1042T did not grow in and AA- 1042*, respectively, contained polyamine submerged cultures, therefore cells were harvested patterns in which spermidine and spermine were the from R agar slant cultures. Polyamine extractions and predominant polyamines. Recently, six species of the analyses were performed as described recently genus Propionibacterium, including the LL-A,pm-con- (Altenburger et al., 1997). taining species Propionibacterium acidipropionici, Propionibacterium acnes, Propionibacterium jensenii, The analysis of the polyamine patterns of coryne- and Propionibacterium lymphophilurn, Propionibacterium nocardioform actinobacteria with LL-A,pm revealed propionicum Propionibacterium thoenii, the presence of at least four different profiles (Table 1). and have been The majority of strains, which belong to the genus shown to contain similar polyamine patterns Nocardioides, were characterized by the presence of (Hamana, 1995). All these six species contained high concentrations of the diamines putrescine and spermidine and spermine as the predominant com- cadaverine. The similarity in the polyamine patterns is pound in the polyamine pattern, demonstrating the close phylogenetic relatedness to the genera in agreement with the phylogenetic homogeneity of the Propioniferax, Luteococcus, Friedmanniella species of the genus Nocardioides (Collins et al., 1994). and Nocardiodes simplex Nocardiodes Microlunatus (Nakamura et al., 1995; Schumann et IMET 10283 and al., et al., et al., plantarum NCIMB 12834Twere found to contain rela- 1997; Tamura 1994; Yokota 1994). Based on the data shown, as well as already published tively low polyamine concentrations and displayed Propionibacterium, only cadaverine as the predominant diamine. The data on species of the genus it appears that members of the family Propioni- relatively high amounts of spermine were in the same bacteriaceae et al., range as found in the other strains of the genus (Stackebrandt 1997) are charac- Nocardioides and it is most likely that this tetraamine terized by the presence of the major compounds is taken up from the growth medium as was previously spermidine and spermine in their polyamine patterns. supposed for members of the genera Clavibacter and Intrasporangium calvum IMET 78 16Tand Terrabacter

180 International Journal of Systematic Bacteriology 49 Polyamine profiles in Actinohacteria

Table 1. Polyamine patterns of coryneform and nocardioform actinomycetes with LL-A,pm as diagnostic diamino acid of their pept idoglyca n

Organism* Growth Polyamine content bol(g dry wt)-'lj medium? DAP PUT CAD SPD HSPD SPM TYR

Intrasporangiunt calvunt I M ET 78 16T ( = 7 KIP"', DSM 43043"') R 0.17 2.02 0.30 1.03 0.05 0.31 0.03 (4.3) (51.7) (7.6) (26.9) (1.3) (7.9) (0.8) Terrabacter tumescens IMET 1043IT (= NCIB 8914"',DSM 2030gT) R 1.35 Tr 0.04 Tr 0.20 (84.9) (2.5) ( 12.6) Terracoccus luteus IMET 784gT (= DSM 44267"') R 0.07 2.04 0.55 1.98 2.16 Tr (1.2) (30.0) (8.1) (29.1) (31.8) Junibacter limosus DSM 11 140T (IF0 16128 = HKI 83T) R 0.07 1.36 0.73 0.06 0.05 0.24 0.09 (2.7) (52.3) (281) (2.3) ( 1'9) (9.2) (3.5) Janibacrer limosus DSM 1 1 141 (IF0 1612gT = HKI 84) R 0.02 1.14 0.84 0.50 - 0.25 0.13 (0.7) (39.6) (29.2) (17.4) (8.7) (4.5) Sanguibacter keddieii DSM 10542T (= ATCC 51767T = NCFB 302ST) R 0.0 1 0.06 0.07 0.12 0.0 1 0.40 Tr (1.5) (9.0) (10.4) (17.9) (1.5) (59.7) Sanguibacter suarezii DSM 10543"'(= NCFB 3023) R 0.03 0.02 0.02 0.05 - 0.31 Tr (7.0) (4.7) (4.7) (1 1.6) (72.1) Nocardioides albus IMET 7807T (= DSM 43109T) R 0.08 5.23 5.16 Tr Tr 0.07 (0.8) (496) (49.0) (0.7) Nocardioides alhus IMET 7807"'(= DSM 43 109=) MM ~ 0.04 0.45 0.02 ~ (7.8) (88.2) (3-9) Nocardioides albus IMET 7801 (= DSM 43364) R 0.08 0.97 2.71 006 Tr 005 Tr (2.1) (25.1) (70.0) (1.6) (1.3) Nocardioides albus IMET 78 15 (= DSM 43365) R 0.14 3.48 3.45 012 - 0.02 0.03 (1.9) (48.1) (47.7) (1.7) (0.3) (0.4)

Nocardioides albus IMET 781 5 (= DSM 43365) MM 0.21 0.78 0.03 ~ (20.6) (76.5) (2.9) Nocardioides albus IMET 7822 (= DSM 43873) R 0.04 3.11 4.59 012 Tr 0.04 (05) (39.4) (58.1) (1.5) (0.5) Nocardioides albus IMET 7835 (= DSM 43874) R 0.23 0.47 0.82 0.02 - Tr Tr (14.9) (30.5) (53.2) (1.3) Nocardioides simplex IMET 1036gT (= NCIB 8929T,DSM 20130"') R 0.20 0.72 2.71 0-89 Tr 0.27 - (4.2) (15.0) (56.6) (18.6) (5.6) Nocardioides siniplex IMET 10283 R 0.16 0.40 0.05 028 0.15 (1 5.4) (38.5) (4.8) (26.9) (14.4) Nocurdioides plantarum NCIMB 12834T (= DSM 11054"') R 0.04 0.20 0.03 - 025 007 (6.8) (33.9) (5.1) (42.4) (1 1.9) Nocardioides jensenii IMET 1067gT (= NCIB 9770T,DSM 2064IT) R 0.07 0.61 1.51 0.34 - 0.26 (2.5) (21.9) (54.1) (12.2) (9.3) Nocardioides lureus IMET 7830T (= DSM 43366"') R 0.15 1.63 3.10 0.65 0.14 0.10 (2.6) (28.2) (53.7) (11.3) (2.4) ( 1.7) Aeromicrohium er.vthreum NRRL B-338IT(= DSM 8599T) R 0.18 0.16 2.22 1.70 0.31 (3.9) (3.5) (48.6) (37.2) (6.8) Aeromicrohium,fastidiosum IMET 7849T (= NCIB 12713T,DSM 10552T) R 0.03 0.1 1 0.25 0.03 - 0.59 Tr (3.0) (109) (24.8) (3.0) (58.4) Sporichthya polvmorpha IMET 7840T (= DSM 461 1 3T) R 0.07 0.48 0.30 2.08 0.04 0.06 0.09 (2.2) (15.4) (9.6) (66.7) (1.3) (1.9) (2.9) Friedmanniella antarcrica AA-1042T(= DSM 11053T) R 0.15 0.26 0.16 2.93 0.03 3.84 0.16 (2.0) (3.5) (2.1) (38.9) (0.3) (51.0) (2.1) Propionijerax innocua NCTC 1 1082T (= DSM 8251"') R Tr 0.21 014 0.77 Tr 0.49 - ( 13.0) (8.7) (47%) (304) Microlunatus phosphovorus JCM 9379T (= DSM 10555T) R 0.91 005 0.96 2.64 0.02 2.19 008 (1 3.3) (0.7) (14.0) (38.5) (0.3) (32.0) (1.2) Luteococcus japonicus IF0 12422T (= DSM 10546T) R 0.05 0.08 0.1 1 0.80 - 0.52 (3.2) (5.1 (7.1) (51.3) (33.3) * DSM, Deutsche Sammlung von Mikroorganismen und Zellkulturen, Braunschweig, Germany ; IFO, Institute for Fermentation, Osaka, Japan ; IMET, Nationale Kulturensammlung fur Mikroorganismen, Zentralinstitut fur Mikrobiologie und Experimentelle Therapie, Jena, Germany (now included in the DSMZ, Braunschweig, Germany) ; JCM, Japan Collection of Microorganisms, Institute of Physical and Chemical Research, Saitama, Japan; NCIB, National Collection of Industrial Bacteria, Aberdeen, UK ; NCFB, National Collection of Food Bacteria, AFRC Institute of Food Research, Reading, UK ; NCIMB, National Collection of Industrial and Marine Bacteria, Torry Research Station, Aberdeen, UK ; NCPPB, National Collection of Plant-pathogenic Bacteria, Central Science Laboratory, York, UK ; NCTC, National Collection of Type Cultures, Central Public Health Laboratory, London, UK ; NRRL, ARS Culture Collection, Northern Regional Research Laboratory, US Department of Agriculture, Peoria, IL, USA. f' Growth conditions as described in the text. $ DAP, 1,3-diaminopropane; PUT, putrescine ; CAD, cadaverine ; SPD, spermidine ; HSPD, sym-homospermidine ; SPM, spermine ; TYR, tyramine; Tr, trace [less than 0.01 pmol (g dry weight)-l]; -, none detected. Values in boldface type are those for major polyamines (which accounted for more than 20% of the total polyamines). Values in parentheses are percentages of the total polyamine content.

lnterna tional Journal of Systematic Bacteriology 49 181 H.-J. Busse and P. Schumann

Table 2. Differential chemotaxonomic characteristicsof coryneform and nocardioform actinomycetes with LL-A,pm as diagnostic diamino acid of their peptidoglycan

Taxon Polyamine Peptidoglycan Diamino acid and Polar lipids1 Major References pattern* interpeptide amino acid at menaquinone(s)!$ bridge position 1 of peptide subunit?

Pvopionibacteriaceae Friedmanniella SPD, SPM GlY LL-A,pm, Gly PI, PG, DPG, PL MK-9(H4) Schumann et al. (1997) Microlunatus SPD, SPM GlY LL-A,pm, Gly PI, PG, DPG, PL MK-9(H4) Nakamura et al. (1995); Schumann et a/. (1997) Luteococcus SPD, SPM GlY LL-A,pm, L-Ala PI, PG, DPG, GL MK-9(H4) Schumann et al. (1 997) ; Tamura et al. (1 994) Prop ioniferax SPD, SPM GlY LL-A,pm, L-Ala PE, PG, PL, GL MK-9(H4) Schumann et al. (1997); Yokota et al. (1994) Nocardioidaceae Aeromicrobium CAD (SPD) Gly LL-A,pm, L-Ala PE, PG MK-9(H4) Miller et al. (1991); Tamura & Yokota (1994) No ca r dio ides CAD (PUT) Gly LL-A,pm, L-Ala PG, DPG, PL, MK-8(H4) Collins et al. (1989); PG-OH O’Donnell et al. (1 982) ; Prauser (1 986) In trasporangiaceae Terrabacter PUT GlY3 LL-A,pm, L-Ala PE, PI, DPG, PL MK-8(H4) Collins et al. (1989) Terracoccus PUT, SPD, Gly, LL-A,pm, L-Ala PE, PI, DPG, PL MK-8(H4) Prauser et al. (1997) SPM Intrasporangium PUT, SPD GlY 3 LL-A,pm, L-Ala PI, PIM, PG, DPG MK-8 Kalakoutskii (1989) ; Schumann et al. (1 997) Janibacter PUT, CAD, None meso-A,pm, none DPG, PG, PI MK-8(H4) Martin et a/. (1997) (SPD) Sanguibacter SPD Ser-D-Glu L-LYS,L-Ala ND MK-9(H4) Fernindez-Garayzabal et al. (1995) Sporickthyaceae Spo r ich t hya SPD GlY LL-A,pm, L-Ala PI, PG, DPG, PL MK-9(H6), Rainey et al. (1993); MK-9(H8) Schumann et al. (1 997) * CAD, cadaverine; PUT, putrescine; SPD, spermidine; SPM, spermine. Polyamines given in parentheses were not detected in all strains of the respective taxon. 7 LL-A,pm, LL-diaminopimelic acid ; rn-A,pm, m-diaminopimelic acid. $+ PI, phosphatidylinositol ; PG, phosphatidylglycerol; DPG, diphosphatidylglycerol ; PL, unknown phospholipid(s) ; GL, unknown glycolipid(s) ; PE, phosphatidylethanolamine ; PG-OH, phosphatidylglycerol containing 2-hydroxy fatty acids. ND,Not determined. !$Abbreviations for menaquinones are examplified by MK-9(H4), menaquinone with two of nine isoprene units hydrogenated.

tumescens IMET 10431 have been grouped together spermidine and spermine were found. Thus the in the family Zntrasporangiaceae (Stackebrandt et al., polyamine pattern of these three organisms does not 1997) and Terracoccus luteus IMET 784ST, which has only display their phylogenetic relatedness but also been described recently (Prauser et al., 1997), can be appears to be useful for their differentiation at the considered as another member of this family. The genus level. However, the genus specificity of the polyamine patterns of these three phylogenetic polyamine patterns has to be confirmed by inclusion of relatives were similar in respect to the presence of more strains of the three genera, which are not yet putrescine as a predominant polyamine. In addition, in available. In order to substantiate the predominant strain IMET 78 16Ta high concentration of spermidine diamine putrescine as a family characteristic feature in was detected and in IMET 7848T high amounts of the polyamine pattern of the Zntrasporangiaceae,

182 lnterna tiona/ Journal of Systematic Bactefio/ogy 49 Polyamine profiles in Actinobacteria

members of two more genera belonging to the family, including Strep tomyces griseus, Strep tomyces Janibacter (Martin et al., 1997) and Sanguibacter chrysomallus, Streptomyces coelicolor, Streptomyces (Fernhndez-Garayzabal et al., 1995) were included in caeruleus, ' Streptomyces thermoolivaceus ', Strepto- our study. Janibacter limosus is known to possess myces thermoviolaceus, Streptomyces thermo- meso-A,pm whereas the two species Sanguibacter diastaticus and Streptomyces thermovulgaris. All these - keddieii and Sanguibacter suarezii contain L-lysine in species contain putrescine as a major pol yamine and, their peptidoglycans. The two strains of J. limosus, in addition, most of the species also contain cadaverine DSM 11140T and DSM 11141, were found to have a and spermidine as major compounds. Due to the polyamine pattern which contained the predominant limited data on the polyamine content of Streptomyces compounds putrescine and cadaverine. In addition, species it cannot be decided whether the polyamine strain DSM 11141 contained significant amounts of approach is applicable for distinction between spermidine. The presence of putrescine in the poly- Sporichthya and Streptomyces. However, the detection amine pattern is in aggreement with the affiliation of of putrescine as a predominant polyamine in Strepto- these two strains with the family Intrasporangiaceae myces species might be a distinguishing characteristic. but the polyamine pattern was clearly distinct from the The polyamine patterns detected in the heterogeneous other members of the family and also appears to be group of coryne- and nocardioform actinobacteria genus-specific. A common characteristic of the poly- amine patterns of the two species S. keddieii DSM with LL-A,~~in the peptidoglycan demonstrated that 10542Tand S. suarezii DSM 10543Twas the relatively this approach is also useful for distinction of the major lines of descent within this group, usually at the family low polyamine content when compared to the other level, and supports a reliable identification at the genus members of the family. In addition, these two species level. Classification and identification of these contained spermine as the predominant compound whereas significant amounts of putrescine were lack- organisms at the genus level necessitates the additional investigation of other chemotaxonomic characteristics ing. This observation demonstrates that putrescine as such as peptidoglycan type, fatty acid and polar lipid a predominant compound is not present in all members profile, and quinone system (Schumann et al., 1997). of the family and thus cannot be considered as a The examination of supplementary characteristics is family-specific characteristic. However, the two necessary, since the polyamine patterns detected for Sanguibacter species are the deepest phylogenetic Actino- et al., LL-A,pm-containing members of the class branching within the family (Prauser 1997). bacteria are not unique within this class. Similar They differ from the other members of the family in patterns are found in other groups of the class respect to their peptidoglycan diamino acid (L-lysine), Actinobacteria, for example the pol yamine pattern their menaquinone MK-9(H4) and their fatty acid Nocardioides J. et al., characteristic for the branch and profile (Prauser 1997). These data, as well as the limosus is similar to that found in Brevibacterium polyamine patterns, confirm the phylogenetic analyses species and the polyamine pattern of the family indicating that the genus Sanguibacter represents a Propionibacteriaceae resembles that found in the gen- separate branch within the In trasporangiaceae era Clavibacter, Rathayibacter and Curtobacterium (Prauser et al., 1997), which might be considered as a (Altenburger et al., 1997; Hamana, 1995). On the subfamily. For pragmatic reasons it would be ap- other hand, the distinguishing polyamine patterns propriate to exclude the genus Sanguibacter from the appear to facilitate differentiation between LL-A,pm- family Intrasporangiaceae and to propose the new containing taxa which have other characteristics in family Sanguibacteriaceae. This transfer would be common, such as the genera Terrabacter and justified by the significant differences in the chem- Nocardioides, both containing MK-8(H4), or the genus otaxonomic characteristics (Table 2). The remainder Aeromicrobium and taxa of the family Propioni- of the family Intrasporangiaceae would be restricted to bacteriaceae, which both contain MK-9(H4) (Table 2). the genera Intrasporangium, Terrabacter, Terracoccus Thus, the polyamine approach is considered as an and Janibacter. important method for classification of coryne- and nocardioform LL-A,pm-containing actinobacteria The only representative of a separate line of descent, when applied in combination with other, well-estab- Sporichthya polymorpha IMET 7840T, could be dis- lished chemotaxonomic investigations. tinguished from the other LL-A,pm-containing taxa by its distinct polyamine pattern. It was characterized by spermidine as the predominant compound and smaller Acknowledgements amounts of putrescine and cadaverine. As long as no We are grateful to Eva Woitzik and Carmen Schult for more strains of this line of descent are available, the excellent technical assistance and to Oliver Kim for critical significance of this polyamine pattern cannot be reading of the manuscript. evaluated. In respect to the major menaquinone and pep tidoglycan characteristics, Sporich thya polymorpha cannot be distinguished from the genus Streptomyces References (Schumann et al., 1997). So far, only eight species of Altenburger, P., Klmpfer, P., Akimov, V. N., Lubitz, W. & Busse, the genus Streptomyces have been analysed for their H.-J. (1997). Polyamine distribution in actinomycetes with group polyamine content (Hamana & Matsuzaki, 1987) B peptidoglycan and species of the genera Brevibacterium,

International Journal of Systematic Bacteriology 49 183 H.-J. Busse and P. Schumann

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