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Peptidoglycan .Types of Bacterial Cell Walls and their Taxonomic ...

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446<br />

depicted in Fig. 24b. The interpeptide bridge<br />

consists <strong>of</strong> fl-D-Asp-L-Ala.<br />

Strain ATCC 11859, called B. pasteurii,<br />

shows the same peptidoglycan type as B.<br />

sphaericus <strong>and</strong> is most likely a urease-producing<br />

B. sphaericus. A third strain (own isolate)<br />

contains a fl-DAsp-L-Ser interpeptide bridge.<br />

Some <strong>of</strong> the L-Ser residues are replaced by<br />

L-Ala. Although more strains <strong>of</strong> both species<br />

need to be investigated, the peptidoglycan<br />

type is most likely a good criterion to distinguish<br />

B. pasteurii from B. sphaericus.<br />

Genus Clostridium. The cell wall composition<br />

<strong>of</strong> these anaerobic spore-forming organisms<br />

<strong>of</strong> the genus Clostridium was studied primarily<br />

by Cummins <strong>and</strong> co-workers (61, 73, 77). They<br />

determined qualitatively the sugar <strong>and</strong> amino<br />

acid composition <strong>of</strong> the cell walls <strong>of</strong> various<br />

clostridia. There are also a few studies on the<br />

quantitative amino acid composition (221, 296,<br />

376). All the available data on the amino acid<br />

composition <strong>of</strong> the cell walls <strong>of</strong> clostridia are<br />

compiled in Table 30.<br />

Most <strong>of</strong> the species investigated contain only<br />

m-Dpm, Ala, <strong>and</strong> Glu. Unpublished studies<br />

from our laboratory on cell walls <strong>of</strong> C. butyricum<br />

have shown that the peptidoglycan <strong>of</strong> this<br />

organism belongs to the directly cross-linked,<br />

m-Dpm-containing type (Aly). The other m-<br />

Dpm-containing peptidoglycans <strong>of</strong> the genus<br />

Clostridium presumably show the same type.<br />

Four different species reveal (instead <strong>of</strong> m-<br />

Dpm) L, L-Dpm <strong>and</strong>, in addition, Gly in <strong>their</strong><br />

cell walls. The primary structure <strong>of</strong> the peptidoglycan<br />

<strong>of</strong> C. perfringens was established by<br />

Leyh-Bouille et al. (221). This species shows<br />

the same peptidoglycan type as that described<br />

before for Propionibacterium petersonii <strong>and</strong><br />

some Streptomyces sp. (Fig. 15a). The three<br />

other species C. fallax, C. pectinovorum, <strong>and</strong> C.<br />

pseud<strong>of</strong>allax probably belong to the same type.<br />

The species C. innocuum, C. paraputrificum,<br />

<strong>and</strong> C. tertium contain L-Lys instead <strong>of</strong> Dpm<br />

in <strong>their</strong> vegetative cell walls. Since no detailed<br />

studies are available, the peptidoglycan type <strong>of</strong><br />

these species is unknown. C. innocuum has<br />

probably been misclassified in the genus Clostridium,<br />

since it differs not only in the amino<br />

acid composition <strong>of</strong> its cell walls but also in its<br />

GC content (43%) from the other strains <strong>of</strong><br />

clostridia (GC content, 22-28%) (77).<br />

The status <strong>of</strong> the other clostridia which<br />

contain no Dpm is uncertain despite <strong>their</strong><br />

rather low GC content (24-27%) which is in<br />

good agreement with that <strong>of</strong> the other members<br />

<strong>of</strong> the genus clostridia. Further studies are<br />

necessary to clarify the taxonomic position <strong>of</strong><br />

these organisms.<br />

SCHLEIFER ANJD<br />

KANDLER BACTERIOL REV.<br />

The peptidoglycans within the genus Clostridium,<br />

as in the genus Bacillus, are rather<br />

uniform. Most <strong>of</strong> the clostridia contain the<br />

directly cross-linked m-Dpm peptidoglycan<br />

type (variation Aly), but a few species are<br />

distinguished by the occurrence <strong>of</strong> the<br />

L, L-Dpm-Gly type. The occurrence <strong>of</strong> other<br />

peptidoglycan types in strains called Clostridium<br />

can be taken as an indication that these<br />

organisms may be improperly classified.<br />

The determination <strong>of</strong> the structure <strong>of</strong> the<br />

polysaccharides <strong>of</strong> the cell walls <strong>of</strong> clostridia<br />

may be a useful tool for characterization <strong>of</strong><br />

species <strong>and</strong> infraspecific taxa within the genus.<br />

Cummins <strong>and</strong> Johnson (77) had already<br />

achieved some separation by means <strong>of</strong> a qualitative<br />

determination <strong>of</strong> the sugar composition<br />

in the cell walls.<br />

Family Corynebacteriaceae. The family<br />

Corynebacteriaceae is characterized by irregular<br />

rod-shaped cells which usually occur in<br />

angular or palisade formation due to snapping<br />

division. Moreover, a marked diversity in morphology<br />

<strong>and</strong> a vegetative life cycle are <strong>of</strong>ten<br />

found. According to the 7th edition <strong>of</strong> Bergey's<br />

Manual (49), the family Corynebacteriaceae<br />

comprises six different genera: Corynebacterium,<br />

Arthrobacter, Microbacterium, <strong>Cell</strong>ulomonas,<br />

Listeria, <strong>and</strong> Erysipelothrix. More<br />

recent studies have shown that almost all <strong>of</strong> the<br />

organisms included in the family Brevibacteriaceae<br />

are also typical coryneform bacteria<br />

(161, 264, 332; Cziharz, Diplom Thesis, Technical<br />

University, Munich, 1969). Therefore, the<br />

organisms classified in the family Brevibacteriaceae<br />

(genera Brevibacterium <strong>and</strong> Kurthia)<br />

will be discussed together with the Corynebacteriaceae.<br />

The lack <strong>of</strong> proper criteria for differentiation<br />

makes the classification <strong>of</strong> most <strong>of</strong> the coryneform<br />

organisms rather unsatisfactory. In the<br />

-G-M-G-<br />

L- Ala<br />

D-Glu NH2<br />

I Y E T<br />

L-Lys e D-Asp - a<br />

4 - E L-Ala--- D-Asp- b<br />

D- Ala<br />

D-Ata<br />

I L-Lys<br />

FIG. 24. Fragments <strong>of</strong> the primary structures <strong>of</strong><br />

uncommon peptidoglycans in the genus Bacillus. (a)<br />

L-Lys-D-Asp, B. sphaericus; (b) L-Lys-L-Ala-D-Asp,<br />

B. pasteurii.<br />

I<br />

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