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Fundamental Food Microbiology, Third Edition - Fuad Fathir

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178 FUNDAMENTAL FOOD MICROBIOLOGY<br />

make fermented sausages, such as phosphate or nitrite, and sanitizers from equipment.<br />

These factors can prevent or reduce the growth of starters.<br />

E. Bacteriophages of Lactic Acid Bacteria<br />

The role of bacteriophages (or phages) in starter-culture failure in food fermentation<br />

has been recognized for a long time. A short discussion on their life cycle that results<br />

in starter failure, as well as some current practices used to overcome phage problems<br />

in food fermentation, is presented here. 6–10<br />

1. Morphology and Characteristics<br />

Bacteriophages are filterable viruses of bacteria widely distributed in the environment,<br />

especially in food fermentation environments. A phage contains several proteins<br />

(that make the head, tail, tail fiber, and contractile sheath) and DNA, which<br />

can be linear or circular, double stranded of ca. 20 to 55 kb in size. The doublestranded<br />

DNA molecule is packed in the head, which can be round or hexagonal<br />

(prolate or isometric, respectively).<br />

Several schemes have been proposed at different times to classify the bacteriophages<br />

of lactic acid bacteria. Currently, a scheme is used in which the family,<br />

group, and morphotype are classified together. The three families Myoviridae,<br />

Siphoviridae, and Podoviridae are included in three groups, A (with contractile<br />

tails), B (with noncontractile long tails), and C (with noncontractile short tails),<br />

respectively. They can have three morphotypes, namely 1 (with small isometric<br />

heads), 2 (with small prolate heads), and 3 (with large prolate heads), respectively.<br />

Lac. lactis ssp. lactis and ssp. cremoris appear to have many wide varieties of<br />

phages. They have been divided into 12 species, of which 3 are in the Siphoviridae<br />

family and are important because of their virulent nature. The 3 in the Siphoviridae<br />

family are 936 species (all are virulent with small isometric head; e.g., sk1), c2<br />

species (all are virulent with prolate head; e.g., c2), and P335 species (have both<br />

virulent and temperate phages and small isometric heads; e.g., TP901-1). Genome<br />

sequence analysis indicates that the species in the same family as well as those<br />

in different families vary considerably. In contrast, the genome analysis of phages<br />

of Streptococcus thermophilus (e.g., sfi 19, sfi 11) indicates a fairly good homology.<br />

Phages of several species of Lactobacillus have been identified and genome<br />

sequences of some species and strains are now available, namely for Lactobacillus<br />

delbrueckii (LL-H), Lab. plantarum (phi-gle), Lactobacillus johnsonii (Lj 965),<br />

Lab. gasseri (adh), and Lab. casei (a2). Very few phages of Leuconostoc and<br />

Oenococcus have been isolated, and none have been isolated from Pediococcus. 6<br />

2. Life Cycle<br />

A phage cannot multiply by itself in food. Instead, it attaches (needs Ca 2+ for<br />

adsorption) with its tail on the surface of a bacterial cell (specific host) and injects<br />

its DNA inside the cell cytoplasm. If it is a lytic phage, the bacterial cell produces<br />

a large number of copies of the phage DNA and phage proteins. Following

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