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

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442 SCHLEIFER AND KANDLER<br />

L-Ala, L-Ala-L-Ala, or L-Ala. The amino acid<br />

compositions <strong>of</strong> the interpeptide bridges <strong>of</strong> the<br />

two subspecies <strong>of</strong> L. viridescens (170) are quite<br />

similar but the amino acid sequences are reversed<br />

(Fig. 21a <strong>and</strong> b). rhus the differences in<br />

the peptidoglycan structure confirm the separation<br />

<strong>of</strong> these two subspecies which was originally<br />

based on physiological <strong>and</strong> morphological<br />

evidence.<br />

The two subspecies <strong>of</strong> L. coprophilus are<br />

distinguished by the number <strong>of</strong> alanine residues<br />

in the interpeptide bridge <strong>and</strong> by the ability <strong>of</strong><br />

one to form dextran (144). Although the peptidoglycan<br />

type is quite similar to that <strong>of</strong><br />

Leuconostoc (vide infra), both subspecies belong<br />

to the subgenus Betabacterium since they<br />

produce D, L-lactic acid <strong>and</strong> split arginine.<br />

L. coprophilus <strong>and</strong> L. viridescens form, in<br />

contrast to other species <strong>of</strong> the genus Betabacterium,<br />

irregular rods, <strong>and</strong> in particular both<br />

subspecies <strong>of</strong> L. coprophilus, <strong>and</strong> L. viridescens<br />

ssp. minor can easily be confused with Leuconostoc.<br />

They may constitute an intermediary<br />

group between the genera Leuconostoc <strong>and</strong><br />

Betabacterium. Williams <strong>and</strong> Sadler (422) report<br />

the occurrence <strong>of</strong> Gly in cell walls <strong>of</strong> some<br />

strains <strong>of</strong> heter<strong>of</strong>ermentative lactobacilli.<br />

These cell walls were reinvestigated in our<br />

laboratory by quantitative amino acid analysis<br />

<strong>and</strong> by determining the peptide pattern <strong>of</strong><br />

partial acid hydrolysates. It could be shown<br />

BACTERIOL. REV.<br />

that these cell walls were contaminated by<br />

protein. The real amino acid composition <strong>of</strong> the<br />

peptidoglycan <strong>of</strong> the strains mentioned by<br />

Williams <strong>and</strong> Sadler (422) is in agreement with<br />

the data given in Table 26.<br />

Genus Leuconostoc. The genus Leuconostoc<br />

was classified according to Garvie (104, 106)<br />

<strong>and</strong> Whittenbury (417). The peptidoglycan<br />

types <strong>of</strong> the different species are listed in Table<br />

27. The majority <strong>of</strong> the strains contain a peptidoglycan<br />

<strong>of</strong> the Lys-L-Ser-L-Ala type. The<br />

remainder <strong>of</strong> the strains reveal di-L-Ala, L-Ser-<br />

Ala, or L-Ser-L-Ser peptides as interpeptide<br />

bridges. The correlation <strong>of</strong> the peptidoglycan<br />

type with the present grouping <strong>of</strong> the strains<br />

-G-M-G-<br />

L- Ala<br />

D- Glu --NH2<br />

iYE<br />

L- Lys +- L-Ala(L-Ser)---- L-Ser a<br />

D-Ala<br />

jI* e L-Ser--- L-Alas-L-Ala *--- b<br />

E L-Ala - (L-Ala)-<br />

c<br />

D-Ala<br />

L- Lys<br />

FIG. 21. Fragments <strong>of</strong> the primary structures <strong>of</strong><br />

peptidoglycans <strong>of</strong> lactobacilli <strong>and</strong> leuconostocs. (a)<br />

L-Lys- L-Ala( L-Ser)- L-Ser; (b) L-Lys- L-Ser- L-Ala,;<br />

(C) L-Lys-L-Ala(2).<br />

TABLE 27. Distribution <strong>of</strong> peptidoglycan types within the genus Leuconostoc<br />

Species No. <strong>of</strong> ATCC or NCDOa no. Type Fig. Reference<br />

strains<br />

Leuconostoc cremoris .... 8 NCDO 828 L-Lys-L-Ser-L-Ala2 21b 169<br />

L. lactis ................ 8 NCDO 533, 546 L-Lys-L-Ser-L-Ala2 plus 21b 169<br />

L-Lys-L-Ala2 21c<br />

L. paramesenteroides ssp.<br />

ssp. paramesenteroides 2 NCDO 803, 883 L-Lys-L-Ala2 21c 169<br />

ssp. lactophilum ...... 5 L-Lys-L-Ser-L-Ala2 21b 169<br />

L. gracile (syn. L. oenos) 23 NCDO 1668, 1669, 1674, 1675, L-Lys-L-Ala-L-Ser plus 21a 169<br />

1694, 1696, 1705, 1707, 1823, L-Lys-L-Ser-L-Ser 21a<br />

1890, 1893<br />

L. dextranicum .......1. NCDO 517 L-Lys-L-Ser-L-Ala2 21b 169<br />

L. mesenteroides<br />

ssp. mesenteroides .. 18 NCDO 183, 523, 538; ATCC L-Lys-L-Ser-L-Ala2 21b 169, 174<br />

8082, 8086, 8293, 8357,<br />

8359, 9135, 10830, 10830a,<br />

10878, 12291, 14430, 14935<br />

ssp. amelibiosus .... 4 ATCC 13146, 10882, 10883, L-Lys-L-Ala2 21c 169<br />

11449<br />

a See footnote a in Table 20.<br />

T<br />

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