Brief history of the taxonomy of Listeria<\/u><\/strong><\/p>\n Introduction<\/u><\/strong><\/p>\n Taxonomy is a scientific discipline pertaining to the classification, nomenclature, and identification of organisms. Organisms are classified or grouped based on shared characteristics, previously unknown strains are placed in known taxa and assigned names following set nomenclature standards. The naming of organisms dates back as early as the great philosopher Aristotle (384-322BC). Modern taxonomy is based on the principles established by the Swedish botanist, Carolus Linnaeus (1707-1778). He introduced binomial nomenclature for naming organisms and established the Linnaean classification system whereby organisms are ranked hierarchically at species, genus, family, order, class, phylum and kingdom levels.<\/p>\n In the last few years, the taxonomy of the genus Listeria has undergone extensive modifications, mainly because of the now well-established DNA hybridisation techniques. Murray, Webb and Swann described Listeria monocytogenes<\/em> as the causative agent for sepsis in rodents in 1926 and they named it Bacterium <\/em>monocytogenes<\/em>because the infection was monocytotic (Liu, 2013<\/a>). <\/em>The following year, Pirie isolated a similar bacillus from the infected liver of the African jumping mice, the gerbils and named it Listerella <\/em>hepatolytica <\/em>(McLauchlin, 1987<\/a>). Up Until 1940, there was considerable confusion in the nomenclature of L. monocytogenes. J. H. Pirie (1940)<\/a> chose Listerella<\/em> as the generic name in honour of Lord Lister, the British surgeon who pioneered antiseptic surgery. However, this name had already been applied by Jahn in 1906 to a group of slime moulds (Mycetozoa<\/em>) (J. H. Pirie, 1940<\/a>). The generic name Bacterium<\/em> as applied by Murray and collaborators was undesirable because the organism does not possess the characteristics of this genus. The resolution of the Committee on Nomenclature, Third International Congress for Microbiology, New York, 1939, was that in all duplications of generic names, only the one first applied should be considered valid, invalidating the generic name proposed by Pirie, and thus he suggested the name Listeria<\/em> in 1940 (J. H. H. Pirie, 1940<\/a>). Listeria<\/em> was adopted in the sixth edition of Bergey\u2019s Manual of Determinative Bacteriology and approved by the Judicial Commission on Bacteriological Nomenclature and Taxonomy, and it is now the official genus name (Rocourt & Buchrieser, 2007<\/a>).<\/p>\n Listeria<\/em> species known to date are as follows: L. monocytogenes<\/em> (J. H. Pirie, 1940<\/a>), Listeria <\/em>grayi<\/em>, Listeria<\/em> innocua<\/em>, Listeria <\/em>welshimeri<\/em>, Listeria <\/em>seeligeri<\/em>, Listeria <\/em>ivanovii<\/em> (Seeliger, Rocourt, Schrettenbrunner, Grimont, & Jones, 1984<\/a>), Listeria <\/em>marthii<\/em> (Graves et al., 2010<\/a>), Listeria <\/em>rocourtiae<\/em> (Leclercq et al., 2010<\/a>), Listeria <\/em>fleischmannii<\/em> (Bertsch et al., 2013<\/a>), Listeria <\/em>weihenstephanensis<\/em> (Lang Halter, Neuhaus, & Scherer, 2013<\/a>), Listeria <\/em>floridensis<\/em>, Listeria <\/em>aquatica<\/em>, Listeria <\/em>cornellensis<\/em>, Listeria <\/em>riparia<\/em> and Listeria <\/em>grandensis<\/em> (den Bakker et al., 2014<\/a>) and the recently published Listeria <\/em>booriae<\/em> and Listeria <\/em>newyorkensis<\/em> (Weller, Andrus, Wiedmann, & den Bakker, 2015<\/a>). Additionally, two subspecies have been identified within L. <\/em>ivanovii<\/em> [subsp. ivanovii<\/em> and subsp. londoniensis<\/em>, (Boerlin et al, 1992 cited in Weller et al. (2015)<\/a>], L<\/em>. grayi<\/em> [subsp. grayi<\/em> and subsp. murrayi<\/em>(Stuart & Welshimer, 1974<\/a>)], and L. <\/em>fleischmannii<\/em> [subsp. fleischmannii<\/em> and subsp. coloradonensis<\/em>, (den Bakker, Manuel, Fortes, Wiedmann, & Nightingale, 2013<\/a>)].<\/p>\n Listeria<\/em> only had one species, L. monocytogenes<\/em>, for many years. It was only in 1948 that Listeria <\/em>denitrificans<\/em> was added, followed by L. <\/em>grayi<\/em> in 1966, L. <\/em>murrayi<\/em> in 1971, L. <\/em>innocua<\/em> in 1981, L. <\/em>welshimeri<\/em> and L<\/em>. seeligeri<\/em>in 1983 and L. <\/em>ivanovii<\/em> in 1985. The newer members of the species were all added post-2009. The species are named after famous scientists in the sciences field who were mainly involved in microbiology related studies. The newer species are mostly named after the places they were first isolated.<\/p>\n Diversity<\/u><\/strong><\/p>\n Listeria<\/em> species are ubiquitous, being found in most natural and urban environmental habitats. The wide distribution in the environment of the species isolated before 1985 is well documented. For example, the newly isolated species (2009 onwards) were from different environments as follows: L.<\/em> weihenstephanensis <\/em>was isolated from vegetation in a pond in Germany (Lang Halter et al., 2013). L.<\/em> rocourtiae<\/em>was isolated from processed lettuce in Austria (Leclercq et al., 2010). L. booriae<\/em> and L.<\/em> newyorkensis<\/em> were isolated from food processing environments in the USA (Weller et al., 2015). L.<\/em> floridensis, L. aquatica, L. cornellensis, L. riparia<\/em> and L. grandensis<\/em> were isolated from natural and agricultural environments in the USA (den Bakker et al., 2014). L. <\/em>fleischmannii<\/em> was isolated from cheeses in Switzerland and Germany (Bertsch et al., 2013). L. <\/em>marthii<\/em> was isolated from different natural environments in the USA (Graves et al., 2010). This clearly shows that the Listeria species is adapted to various environmental conditions and not confined to specific environments.<\/p>\n The genus Listeria<\/em> is composed of short (0.5 \u2013 2 \u00b5m long), Gram-positive, non-sporulating, facultatively anaerobic rods that have a low G + C content. Cells are non-encapsulated exhibiting growth at temperatures ranging from 0 \u2013 45o<\/sup>C, and at pH 6 \u2013 9. Depending on the growing conditions, the cells can appear coccoid and motile. Listeria <\/em>species are catalase-positive and oxidative-negative. According to Schleifer & Kandler, 1972, cited in Lang Halter et al. (2013)<\/a>, the cell walls contain meso-diaminopimelic acid variation A1\u03b3. When grown at 37o<\/sup>C, the major fatty acids are anteiso-C17:0<\/sub> and anteiso-C15:0<\/sub>. Menaquinones are the sole respiratory quinones as stated by Ludwig et al cited in Lang Halter et al. (2013)<\/a>.<\/p>\n \u00a0<\/strong><\/p>\n Pathogenicity<\/u><\/strong><\/p>\n Three of the Listeria <\/em>species, that is, L. monocytogenes<\/em>, L. ivanovii<\/em> and L. <\/em>seeligeri<\/em> are haemolytic and harbour the Listeria <\/em>virulence gene cluster. Of these species, only L. monocytogenes<\/em> and L. ivanovii<\/em> are pathogenic to animals and humans, with L<\/em>. monocytogenes <\/em>expressing pathogenicity in humans and L. ivanovii<\/em> in ungulates (cattle and sheep). L. <\/em>seeligeri<\/em> is not pathogenic to humans even though it contains homologues of the virulence gene cluster (Lang Halter et al., 2013<\/a>). Although L. monocytogenes<\/em> is recognized as the causative agent of listeriosis in humans, rare cases of infection by L. <\/em>innocua<\/em>, L. ivanovii<\/em>, and L. <\/em>seeligeri<\/em> have also been reported (Magalh\u00e3es et al., 2014).<\/p>\n Batz (2005), Gillespie (2010) and Painter (2013) cited in Weller et al. (2015)<\/a> stated that L. monocytogenes<\/em> poses a significant threat to public health and Scallan et al (2011) cited in Orsi and Wiedmann (2016)<\/a> mentioned that it is the third leading cause of foodborne deaths related to microbial contamination. It is the most commonly isolated member responsible for listeriosis (Leclercq et al., 2010<\/a>). Major outbreaks of food-borne listeriosis have been recorded in different parts of the industrialised world. This has raised grave concerns in relation to public health. L. monocytogenes<\/em> has been identified as the causative agent in all cases of human listeriosis. It mainly manifests in high-risk segments of the populations such as foetuses, neonates, infants, the elderly, pregnant women and immunocompromised people Farber and Peterkin, 1991 cited in Gormon and Phan-Thanh (1995). The ability of L. monocytogenes<\/em> strains to survive in conditions of temperatures as low as 4o<\/sup>C have made it a bacterium that is significant in the food processing sector especially in ready-to-eat foods.<\/p>\n Phylogeny<\/u><\/strong><\/p>\n The relationship of Listeria<\/em> spp to other bacteria was relatively unclear until the 1970s. It only received recognition in in 1930 when it was incorporated in the tribe Kurthia<\/em> of the Corynebacteriaceae<\/em> family. In 1974, it was unclearly placed with Erysipelothrix <\/em>and Caryophanon<\/em> after the Lactobacillaceae <\/em>family. It was subsequently grouped with Lactobacillus<\/em>, Erysipelothrix<\/em>, Brochothrix<\/em>, Renibacterium<\/em>, Kurthia<\/em> and Caryophanon<\/em> in the section of \u201cregular, non-sporing Gram-positive rods\u201d in Bergey\u2019s Manual of Determinative Bacteriology <\/em>(Rocourt & Buchrieser, 2007<\/a>). <\/em><\/p>\n Studies by Jones (1975) paved way for a broader understanding of the phylogenetic relationship between then known Listeria spp and various other genera. Wilkinson and Jones (1977) and Feresu and Jones (1988) cited in Rocourt and Buchrieser (2007)<\/a> carried out studies that progressively led to a phylogenetic refinement in the positioning of Listeria<\/em>. These works clearly distinguished Listeria<\/em> from other genera such as Erysipelothrix<\/em> and Brochothrix <\/em>thermosphacta<\/em> and its relatedness to Lactobacillus<\/em> and Streptococcus.<\/em> Based on these results, Wilkinson and Jones (1988) cited in Rocourt and Buchrieser (2007)<\/a> suggested that Listeria, Gemella, Brochothrix, Streptococcus<\/em> and Lactobacillus<\/em> be classified in the family Lactobacillaceae.<\/em><\/p>\n Chemotaxonomic methods have shown that Listeria<\/em> spp belong to the low G + C percent DNA content (<55%) group of Gram-positive bacteria. Lipoteichoic acids isolated from L. monocytogenes<\/em> exhibit structural analogies with lipoteichoic acids from other bacteria. Rocourt and Buchrieser (2007)<\/a> suggested that they may be used as taxonomic markers. According to Ruhland and Fiedler (1987) cited in Rocourt and Buchrieser (2007), lipoteichoic acids imply biochemical consistency and their absence in Coryneform bacteria and presence in Bacillus, Staphylococcus, Streptococcus <\/em>and Lactobacillus<\/em> cements the grouping of Listeria<\/em> with the latter. Mara and Michalec (1977) cited in Rocourt and Buchrieser (2007) reported the absence of free mycolic acids in Listeria<\/em>. The presence of menaquinones corroborates relatedness of Listeria<\/em> and Brochothrix<\/em> albeit distance from lactobacilli (Rocourt & Buchrieser, 2007<\/a>).<\/p>\n 16S and 23S rRNA analysis of L. monocytogenes has elucidated the position of Listeria<\/em> with regard to the other genera of Gram-positive bacteria. Collins et al cited in Rocourt and Buchrieser (2007) deduced that Listeria<\/em> is remote from Lactobacillus<\/em> and should not be in the Lactobacillaceae<\/em> family and that the Listeria-Brochothrix<\/em> subline should be a different family, Listeriaceae.<\/em> 23S rRNA sequencing has affirmed the dissimilarities between Lactobacillus<\/em> and Listeria<\/em> and the similarities with Bacillus<\/em> and Staphylococcus <\/em>(<\/em>Rocourt & Buchrieser, 2007<\/em><\/a>)<\/em>.<\/p>\n According to den Bakker et al. (2014)<\/a>, the species of the genus Listeria can be put into four clades by use of 16S rRNA gene and amino acid sequence phylogenies as follows:<\/p>\n They also state that the L. rocourtiae<\/em> and L. grayi<\/em> clades are sister groups based on 16S rRNA gene sequence analysis.<\/p>\n Sensu<\/u><\/strong>stricto<\/u><\/strong> and <\/u><\/strong>sensu<\/u><\/strong>lato<\/u><\/strong><\/p>\n Chiara et al. (2015)<\/a> and Orsi and Wiedmann (2016)<\/a>\u00a0 categorised the species in the genus Listeria into two groups. Listeria<\/em> sensu stricto, comprising L. monocytogenes<\/em>, L. seeligeri, L. marthii, L. ivanovii, L<\/em>. welshimeri<\/em> and L. innocua <\/em>and Listeria<\/em> sensu lato, consisting of L. grayi <\/em>and all the other 10 remaining species.\u00a0 These researchers based their separation on the relatedness of the species to L. monocytogenes<\/em> which was the first Listeria<\/em> species to be classified and its importance to the food processing sector.<\/p>\n\n