TABLE 4.

Examples of some microbial species occurring during chilled storage of meat and their potential spoilage effects.

Bacteria	Growth conditions	Spoilage effects	References
Actinetobacter spp.	Especially present in dairy and seafood products.	Low spoilage potential but can enhanced the growth of other spoilage bacteria by means of quorum sensing.	Pinu, 2016; Ghasemi-Varnamkhasti et al., 2018; Odeyemi et al., 2018; Hahne et al., 2019
Brochothrix spp.	In different gas composition, such as under air, modified atmosphere and vacuum-packaging. More tolerant in oxygen-depleted and CO2-enriched environments.	Sour, acid and cheesy odor.	Koutsoumanis et al., 2008; Nychas et al., 2008; Ercolini et al., 2011; Doulgeraki et al., 2012; Zhao et al., 2015; Mann et al., 2016; Del Blanco et al., 2017; Reid et al., 2017; Mansur et al., 2019
Carnobacterium spp.	In all types of packaging conditions. Predominance in low O2 packaging.	Spoilage effect can vary, producing volatile molecules with low sensory impacts (fruity or fermented odors, …)	Casaburi et al., 2011; Doulgeraki et al., 2012; Pothakos et al., 2015
Lactobacillus spp. (Lb. sakei, Lb. fuchuensis, Lb. plantarum, Lb. curvatus, Lb. algidus, Lb. oligofermentans, …)	In all types of packaging conditions. Predominance with high concentration of CO2.	Severe acidification, emission of off-odor compounds and ropy slime. However, lactic acid bacteria may produce lactic acid, which inhibits the growth of other families of bacteria. And some species can produce bacteriocins.	Kato et al., 2000; Fadda et al., 2010; Doulgeraki et al., 2012; Dalcanton et al., 2013; Nieminen et al., 2015; Pothakos et al., 2015; Zhao et al., 2015; Alvarez-Sieiro et al., 2016; Mann et al., 2016; Woraprayote et al., 2016; Stefanovic et al., 2017; Mansur et al., 2019
Lactococcus spp.	In various types of packaging.	Traditionally they have not been considered as spoilage microorganisms, but the spoilage potential of these bacteria is still scarcely known.	Kato et al., 2000; Doulgeraki et al., 2012; Rahkila et al., 2012; Dalcanton et al., 2013; Pothakos et al., 2014; Zhao et al., 2015; Mann et al., 2016; Mansur et al., 2019
Leuconostoc spp. (Ln. gelidum, Ln. carnosum, Ln. mesenteroides, …)	Under aerobic, vacuum and modified atmosphere packaging. Predominance with high concentration of O2.	Buttery aroma, formation of slime, blowing of packages, green discoloration.	Kato et al., 2000; Doulgeraki et al., 2012; Dalcanton et al., 2013; Nieminen et al., 2015; Pothakos et al., 2015; Zhao et al., 2015; Mann et al., 2016; Mansur et al., 2019
Photobacterium spp.	Under air, vacuum and modified atmosphere packaging. More frequently present in seafood products.	Typically not associated with spoilage of meat. Responsible for reducing TMAO to TMA, off-odor (produce volatile organic compounds) and biogenic amine formation. The mechanism underlying spoilage has not been clarified.	Nieminen et al., 2016; Li et al., 2019
Pseudomonas spp.	In different gas composition, such as under air, modified atmosphere and vacuum-packaging. Predominance under aerobic low temperature. Limitation in the bacterial flora by the presence of CO2 and/or the limitation of O2 in MAP packaging.	Slime, discoloration, off-odor producing.	Koutsoumanis et al., 2008; Nychas et al., 2008; Ercolini et al., 2011; Andritsos et al., 2012; Doulgeraki et al., 2012; Zhao et al., 2015; Mann et al., 2016; Del Blanco et al., 2017; Reid et al., 2017; Liu et al., 2018; Spanu et al., 2018; Mansur et al., 2019
Weissella spp.	Some can be found in salted and fermented foods. Present in vacuum packaging.	Greenish appearance. Can plays an important role in the fermentation process. Some species can produce bacteriocins.	Pothakos et al., 2015; Martins et al., 2016; Kim et al., 2017; Kariyawasam et al., 2019