The aim of this review on lactic acid bacteria has been to indicate what they are (general description), who they are (classification), what they do (metabolism), and how they do it (metabolism, energetics, and transport). This has not been an easy task. LAB comprise a very diverse group of organisms, which have sufficient characteristics in common that some generalizations can be made. The overall view I would like to pass on is that LAB are more than just lactic acid producers. Lactic acid production is merely a reflection of an underlying metabolism, which is far more complex and, most importantly, more adaptive than one could imagine. The results of genome sequencing of LAB will certainly strengthen this view and, to some extent, have already done so. The second LAB genome to be completed, Lb. plantarum strain WCFS1, revealed a large region in the chromosome containing genes encoding several nutrient utilization systems and extracellular functions.
This cluster of genes has been termed a “lifestyle adaptation region”.[19]
LAB are perfectly adapted to environments rich in nutrients and energy sources.
They have, therefore, dispensed with biosynthetic capability. Apparently genetic material for biosynthesis is still present, as shown for the sequenced Lb. plantarum strain mentioned above.[19]However, the nutrient requirements in minimal media[317]are greater than the sequence implicates, indicating that some of the biosynthetic genes are not expressed.
The reasons for this are not known, but early studies showed that mutagenesis can render some lactobacilli prototrophic for some amino acids.[250] However, if these mutants are returned to a rich medium, they readily revert to the auxotrophic state. This may have evol-utionary implications, but more important, it shows that there has been a strong selection for cells that are committed to life in rich environments. These environments are of course excellent for supporting growth of other microorganisms. LAB have therefore developed strategies to efficiently compete with these organisms. One important strategy is acid pro-duction and acid tolerance.[318]This may be why the term LAB arose. A group of bacteria were isolated from similar niches and turned out to be lactic acid producers. It was natural to group them together. Another possibility is the following: the commitment to life in rich environments demands a simple, but effective way of outcompeting other microorganisms.
Solution: acid production! Therefore, we are left with, in reality (phylogenetically), a group of organisms that are diverse but physiologically similar, since they are specialized for nutrient-rich environments (limited biosynthetic ability) and their metabolism is aimed at acid production.
Although the reasoning above may be somewhat oversimplified, it is clear that the classification problems that always have been evident with regard to LAB may stem from an (historical) overemphasis on a few characteristics. We now have the means to determine natural relationships more objectively and more accurately than ever before.
These relationships can probably be assessed more easily than defining common phenoty-pic characters for a particular natural group. In future classification of bacteria, in general,
Hoang-Dung TRAN and friends it will be necessary to do “reverse phylogenetics,” i.e., first define the natural relationships among bacteria with, e.g., rRNA sequencing and then search for the unifying phenotypic characteristics. Interestingly, some approaches in this direction have been taken with regard to LAB. Some enterococci are known to be able to synthesize cytochromes in the presence of heme. The study by Meisel et al.,[35]in which it was shown that a species of Carnobacterium (in the study designated Lb. maltaromicus) can synthesize cyto-chromes in the presence of heme, was done with the phylogenetically close relationship between carnobacteria and enterococci in mind. The authors (correctly) suspected that car-nobacteria could be similar to enterococci because of a phylogenetic relationship.
The general metabolism and physiology of LAB reflect their adaptation to niches rich in nutrients. They have developed very efficient transport systems, which enable them to quickly take up the necessary solutes. Their extreme saccharolytic nature is another example. Genome sequencing has confirmed this picture. Again, the genome of Lb. plantarum can serve as an example as it revealed an impressive 25 complete sugar PTS systems.[19]However, as I have tried to emphasize, LAB have developed systems that will allow them to derive more energy from a rich medium than just carbohydrate.
One of these systems involves the “cofermentations” that have been mentioned several times. By using a substrate, otherwise nonfermentable, as an electron acceptor during carbohydrate fermentation, they indirectly derive some energy from that substrate that otherwise would be lost. Electrogenic precursor/product exchange, e.g., malate and citrate metabolism, essentially serve the same purpose.
As indicated above and in Sec. III, Orla-Jensen’s concept of LAB being a “great natural group” may not be entirely correct. However, the term lactic acid bacteria will probably be used in the foreseeable future, since it is useful to describe a group of organ-isms that have many physiological properties in common and, as a generalization, have similarities in their ecological behavior.
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