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Taxonomic Significance of Phospholipids in Coryneform and Nocardioform Bacteria

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Taxonomic Significance of Phospholipids in Coryneform and Nocardioform Bacteria J. Gen. Appl. Microbiol., 21, 251-261 (1975) TAXONOMIC SIGNIFICANCE OF PHOSPHOLIPIDS IN CORYNEFORM AND NOCARDIOFORM BACTERIA ICHIRO KOMURA, KAZUHIKO YAMADA, SHIN-ICHIRO OTSUKA, AND KAZUO KOMAGATA* Central Research Laboratories, Ajinomoto Co., Inc., Kawasaki 210, Japan and *The Institute of Applied Microbiology , University of Tokyo, Tokyo 113 (Received March 26, 1975) Phospholipid compositions were radiochemically studied on nocardio- form and coryneform bacteria. Phosphatidylethanolamine (PE) was present in the strains of the genera Corynebacterium, Nocardia, and Myco- bacterium. Cardiolipin and phosphatidylglycerol were found in all the genera tested. Sugar-containing phospholipids such as phosphatidyl- inositolmannosides and phosphatidylinositol were present in about half of the genera tested. The genera Corynebacterium was divided into two sub-groups by the phospholipid composition. Corynebacterium strains having a high GC-content exhibited the presence of a large amount of PE and those having a low GC-content a trace amount of PE. Nocardia and Mycobacterium were different from Actinomadura and Oerskovia in the point of phospholipid profile. The phospholipid composition of Nocardia and Mycobacterium was similar to that of the high GC-content group of Corynebacterium. From these findings, the phospholipid composition is considered to be useful for differentiation of coryneform and nocardio- form bacteria. In the course of taxonomic studies on coryneform bacteria, KOMAGATAet al. (1) and YAMADAand KOMAGATA(2-5) divided this group of bacteria into 7 groups including 5 known genera on the basis of the mode of cell division, DNA base composition, principal amino acid in the cell wall, and biochemical and phy- siological characteristics. MORDARSKAet al. (6) reported that the genus Nocardia contained the lipid LCN-A but the genera Actinomadura, Mycobacterium, Oerskovia and Strepto- myces did not. LECHEVALIERet al. (7) found a-branched, ~3-hydroxylated fatty acid in Nocardia-Mycobacterium-Corynebacterium complex, and reported that these 251 252 KOMURA, YAMADA, OTSUKA, and KOMAGATA VOL. 21 I) U E0 b U 0 b 0 E 0 U0 0 U 4. 0 0 0 0 Oa 0 U U z A b 0 0 0 0 0 U b 0 0 0 Oa 0 a U H 1975 Taxonomic Significance of Bacterial Phospholipid 253 C C 0 U -ti F 254 KOMURA, YAMADA, OTSUKA, and KOMAGATA VOL. 21 0 U 4) cd 1975 Taxonomic Significance of Bacterial Phospholipid 255 organisms could be separated into two groups by the structural pattern of these compounds. However, phospholipid compositions of these bacteria have not been investigated to date. The present authors (8) previously reported the phospho- lipid composition of aerobic gram-positive cocci and its taxonomic significance. This paper deals with the phospholipid composition of coryneform and nocardioform bacteria, and discusses its taxonomic significance in these groups of bacteria. MATERIALS AND METHODS Microorganisms. Number and sources of strains employed are shown in Table 1. Twenty-eight strains of coryneform bacteria, 31 strains of nocardioform bacteria, and two strains of Streptomyces were employed. Typical strains of the coryneform bacteria were selected on the basis of YAMADAand KOMAGATA's study (5), and those of nocardioform bacteria were referred to the studies of Fig. 1. Autoradiogram of 32P-phospholipids of Nocardia erythropolis AJ 9126 at different culture ages. 256 KOMURA, YAMADA, OTSUKA, and KOMAGATA VOL. 21 Fig. 2. Deacylated products of phospholipids of Nocardia erythropolis AJ_9126and Escherichia coli AJ 2617. GPE : Glycerylphosphorylethanolamine. GPG : Glycerylphosphorylglycerol. GPGPG : Glycerylphosphorylglycerylphosphorylglycerol. GPI: Glycerylphosphoryl- inositol. GPIM: Glycerylphosphorylinositolmannosides. Fig. 3. Autoradiogram of 32P-phospholids of Streptomyces strains. 1975 Taxonomic Significance of Bacterial Phospholipid 257 LECHEVALIERet al. (7) and of GOODFELLOW(9). The grouping of coryneform bacteria was followed by YAMADAand KoMAGATA'sstudy, but names of species employed in this work were those used in the culture collections. Analytical methods. Phospholipids were analyzed by using radioactive phos- phoric acid as previously reported (8). DNA base composition. DNA base composition (GC-content in DNA) was determined by the method previously reported by YAMADAand KoMAGATA(3). RESULTS Identification of phospholipids The lipids of Nocardia erythropolis AJ 9126 extracted by SHIBUYAand MARUO's method (8,10) showed five spots of phospholipids on the autoradiogram as shown in Fig. 1. In the solvent system employed in this study, the phospholipids of Escherichia coli were separated in the order of phosphatidylethanolamine (PE), cardiolipin (CL), and phosphatidylglycerol (PG) (8,11). By comparison of Rf values of the phospholipids of E. coli, spots 1, 2, and 3 of N. erythropolis were considered to be PE, CL, and PG, respectively. TAKINAMIet al. (12) and Ko- BAYASHIet al. (13) reported that Corynebacterium hydrocarboclastus, which was reidentified later as N. erythropolis by KOMURAet al. (14), contained PE, CL, phosphatidylinositol (PI), and phosphatidylinositolmannosides (PIM), and that the amount of PI was smaller than that of PIM. As shown in Fig. 1, radioactivity of spot 4 was higher than that of spot 5, and we tentatively presumed spots 4 and 5 as PIM and PI, respectively. The spots of deacylated phospholipids of N. erythropolis AJ 9126 on the auto- radiogram are shown in Fig. 2. From the Rf value in the present solvent sys- tem (15), spots 1, 2, 3, and 4 corresponded to glycerylphosphorylethanolamine, glycerylphosphorylglycerol, glycerylphosphorylglycerylphosphorylglycerol, and glycerylphosphorylinositol, which were derived from PE, PG, CL, and PI, re- spectively. The original radioactive phospholipids were confirmed through these results. Effect of cultural conditions A comparison of phospholipid composition in different growth phases of N. erythropolis AJ 9126 is shown in Fig. 1. Any qualitative differences were not found in the phospholipid composition between the cells grown for 16 and 48 hr. This indicates that the phospholipid composition does not change at different growth phases. It was found that phospholipid composition did not change in the cells cultured in nutrient broth (Difco) and brain heart infusion broth (Nissui, Tokyo). Further, phospholipid compositions of Corynebacterium diphtheriae AJ 1414, Brevibacterium linens AJ 1520, Nocardia asteroides AJ 9128, and Nocardia caviae AJ 9172 also did not change by cultural conditions. 258 KOMURA, YAMADA, OTSUKA, and KOMAGATA VOL. 21 Phospholipid compositions of coryneform and nocardioform bacteria The phospholipid compositions of coryneform and nocardioform bacteria are shown in Table 1. Phosphatidylethanolamine was found in Corynebacterium, Nocardia, and Mycobacterium, and CL and PG were found in all the genera tested. Sugar-containing phospholipids such as PIM and PI were found in about half of the genera tested. The genus Corynebacterium was divided into two sub-groups by the phospholipid composition. Corynebacterium strains having a high GC- content exhibited the presence of a large amount of PE, but those having a low GC-content a trace amount of PE. As 13 strains of N. erythropolis showed the same phospholipid profile, this property is of interest from the species level. The phospholipid composition of Nocardia and Mycobacterium was similar to that of the high GC-content group of Corynebacterium. Further, Nocardia and Mycobacterium were different from Actinomadura and Oerskovia in the point of the phospholipid profile. Phospholipid composition of myceliumforming actinomycetes Phospholipid compositions of mycelium-forming actinomycetes, Streptomyces rimosus AJ 9153 and Streptomyces venezuelae AJ 9154, are shown in Fig. 3. Seven to eight kinds of phospholipid were recognized in these Streptomyces strains, but the phospholipids could not be identified due to insufficient separation of the spots on the autoradiogram. The phospholipid composition of Streptomyces was clearly different from those of coryneform and nocardioform bacteria. DISCUSSION In coryneform bacteria, a large amount of PE was characteristically found in the high GC-content group of Corynebacterium, and a trace amount in the low GC-content group of Corynebacterium. Corynebacterium can be distinguished from other genera of this group of bacteria on the basis of phospholipid com- position. Further, the strains of Nocardia and Mycobacterium showed the presence of PE, and they seem to be related to Corynebacterium on their phospholipid com- position. It is of interest to compare these 3 genera because they have meso- diaminopimelic acid (meso-DAP) in the cell walls in common. The composition of phospholipid profile and other characteristics of coryneform and nocardioform bacteria containing meso-DAP in the cell wall are presented in Table 2. From these characteristics and phospholipid profile, Nocardia and Mycobacterium are considered to be related to the high GC-content group of Corynebacterium, and these genera and the group seemed to be different from Brevibacterium, Actino- madura, and the low GC-content group of Corynebacterium. Nocardia and the high GC-content group of Corynebacterium were similar to each other with the exception of the type of cell division. However, fragmentation which has been recognized in Nocardia may be regarded as multi-snapping, since a zigzag arrangement of daughter cells results from it. In fact, JICINSKA(16) 1975 Taxonomic Significance of Bacterial Phospholipid 259 0 v ~ro 52 o ,o`~ .~ U Q N ,.fl
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