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Page 10 of 15 Kamilari et al. Microbiome Res Rep 2025;4:3 https://dx.doi.org/10.20517/mrr.2024.47

Lactobacillus and Leuconostoc as the dominant taxa in Cypriot sausages and indicated that Leuconostoc was
[1]
mostly associated with industrial, compared to traditional Pitsilia sausages . A greater representation of
Lactobacillus species in Mediterranean fermented meat products, compared to those from Southern Europe,
[4]
was detected by Van Reckem et al. . Furthermore, an elevated relative abundance of Lactobacillus species
was detected in Italian and Spanish spontaneously fermented sausages [22,38,40] . Lactobacilli are among the
[41]
most desired microbes to guide the fermentation process . Their capability to ferment sugars and produce
lactic acid creates an acidic environment essential for fibrillar protein coagulation. This process favors the
[42]
stability and cohesiveness of sausages, facilitating slicing . Additionally, lactobacilli affect the sensory
characteristics of sausages by metabolizing acetic acid, succinic acid, and formic acid . Importantly, they
[43]
are considered gatekeepers against the growth of pathogenic and spoilage bacteria due to the formation of
[44]
organic and other antimicrobial compounds, such as diacetyl, hydrogen peroxide, and bacteriocins . For
instance, bacteriocins such as plantaricin (from Lactiplantibacillus plantarum), or Sakacin (from
Latilactobacillus sakei) have been identified to possess biopreservation properties [45,46] .

Apart from Lactobacillus and Leuconostoc, Lactococcus was detected in high relative abundances in
traditional Mytilini sausages. These bacteria can produce an increased amount of organic acids, including
lactic and acetic, throughout their exponential growth, which, in combination with a conversion to ketone
and alcohols, contribute to the richer sensory characteristics of the produced sausages [47,48] . Importantly,
species from these genera are applied as starters for the food industry, owing to their capacity to generate
antimicrobial peptides that can inhibit the growth of spoilage and pathogenic microbes. According to online
bacteriocin databases Bactibase, Labiocin, and Bagel4, Lactococcus has been reported to produce 40 different
bacteriocins (http://bactibase.hammamilab.org/main.php; https://labiocin.univ-lille.fr/; http://bagel4.
molgenrug.nl/), such as nisin, lacticin, lactococcin G, etc.

Pediococcus sp. was also detected in several Cypriot sausages. The presence of species of this genus, such as
P. acidilactici and P. pentosaceus is considered beneficial since some strains are classified as probiotic .
[49]
Therefore, some strains have been applied as starter cultures in fermented sausages to guide fermentation
[50]
process and acidification degree . In addition, specific strains possess antimicrobial activity against
pathogens due to the formation of pediocin, a bacteriocin with potent antilisterial activity . Pediococcus sp.
[51]
has been identified to create negative associations with molds, such as Stemphylium, Kluyveromyces
marxianus, and Kazachstania barnettii. Ilavenil et al. indicated that a Pediococcus strain isolated from Italian
ryegrass, P. pentosaceus KCC-23, showed effective antifungal activity against species such as Botrytis sp. and
[52]
Fusarium sp. . Additionally, Fugaban et al. isolated two Pediococcus species, P. acidilactici and
P. pentosaceus, with inhibitory activity against several mold species, including Alternaria alternate . In our
[53]
analysis, we revealed a negative association of Pediococcus with spoilage bacteria, such as
Enterobacteriaceae, Erysipelotrichaceae, Clostridium, including C. perfringens, and Pseudomonas spp.,
[54]
members of which are considered foodborne and or opportunistic pathogens .

Members of the genus Clostridium have been detected in industrially produced sausages. Members of the
genus Clostridium, such as C. perfringens, C. botulinum, and C. difficile, are considered important foodborne
pathogens [55-57] . These species produce heat-resistant spores, which can germinate and proliferate in
sausages, producing enterotoxins. The presence of these enterotoxins may lead to gastrointestinal
symptoms, such as diarrhea, or even death [55-57] . Apart from Clostridia, industrially produced sausages
indicate the existence of members of the family Enterobacteriaceae. Their occurrence in sausages is
considered undesirable, due to the production of unwanted flavors and chemical compounds that lead to
[58]
the deterioration or spoilage of the product . Still, they are commonly detected in fresh and frozen meat
products [59,60] . The presence of Clostridia and Enterobacteriaceae in industrially produced sausages indicates