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Aeromonas ME Martino, Institute of Functional Genomics (IGFL), Lyon, France L Fasolato and B Cardazzo, University of Padova, Legnaro, Italy

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Aeromonas ME Martino, Institute of Functional Genomics (IGFL), Lyon, France L Fasolato and B Cardazzo, University of Padova, Legnaro, Italy Aeromonas ME Martino, Institute of Functional Genomics (IGFL), Lyon, France L Fasolato and B Cardazzo, University of Padova, Legnaro, Italy ã 2016 Elsevier Ltd. All rights reserved. Introduction Aeromonas and Laboratory Identification The genus Aeromonas belongs to the Aeromonadaceae family Aeromonas spp. can be easily isolated from clinical and envi- and comprises gram-negative, non-spore-forming, rod-shaped, ronmental samples. Several media are routinely used for Aero- facultative anaerobic bacteria that can be isolated from a very monas isolation, but their performances can vary according to wide spectrum of environmental niches. The history and the the nature of samples (food, clinical, or water) and some perception of Aeromonas by the scientific community have selective agents that can reduce the recovery of some species. evolved over 100 years, from its discovery in the late Aeromonas grow well on routine enteric isolation media nineteenth and early twentieth centuries until nowadays. Aero- (MacConkey, XLD, HE, SS, and DC media); however, the monads were first described as pathogens of poikilothermic lactose-negative isolates must be differentiated from com- animals. Today, they are recognized as causing severe illnesses monly isolated pathogens such as Salmonella and Shigella. in aquatic organisms (fish and other cold-blooded species) and The media that are frequently used for both qualitative and also as emerging pathogens associated with several human quantitative evaluations of Aeromonas spp. are listed in Table 2. infections and, in particular, as food-borne pathogens. In Microbiological methods are clearly needed for bacterial 2010, Janda and Abbott published an excellent review about isolation, but their use as species identification tools can be Aeromonas spp., providing a wide and comprehensive view of very challenging, especially for aeromonads. For instance, it the genus. However, many open questions regarding the ecol- can be difficult to separate A. veronii bv. sobria or A. caviae from ogy, pathogenicity, and taxonomy of aeromonads were A. hydrophila or they may be confused with other genera, such present, and after 4 years, Aeromonas still represents a very as Vibrio and Plesiomonas. complex genus. In the last decade, DNA-based molecular methods have A distinctive characteristic of Aeromonas has always been its become more popular and widely acceptable for bacteria spe- controversial taxonomy. The complexity in identifying and cies identification due to their reproducibility, simplicity, and discriminating Aeromonas species relies on the extremely high high discriminatory power. Several molecular methods have intra- and interspecies genetic variability. The genus was first been applied for discriminating Aeromonas species. 16S rRNA discovered in 1891 and included in the family of Vibrionaceae gene sequencing represents the most commonly utilized together with Vibrio spp., Plesiomonas spp., and Photobacterium molecular technique for this purpose. However, it is now spp. This was due to the prevalence of these genera in the recognized to be problematic for bacterial characterization aquatic environments and the common phenotypic character- mainly because of its intragenomic heterogeneity. This sug- istics. The genus was officially created in 1943 and aeromonads gested that a single-gene-based identification approach may were roughly divided into two major groups, based upon not be appropriate for characterizing Aeromonas spp. As a con- growth characteristics and other biochemical features. The sequence, the multilocus sequence typing (MLST) approach mesophilic group, named A. hydrophila, consisted of motile became the new trend in the last 10 years. From 2011, three isolates that grew well at 35–37 C and were associated with MLST schemes were published for Aeromonas spp. demonstrat- a variety of human infections. The psychrophilic group, ing the validity of this technique in discriminating aeromonads referred to as A. salmonicida, included nonmotile strains that at species level. Moreover, the first Aeromonas MLST online had optimal growth temperatures of 22–25 C and caused database was opened (www.pubmlst.org/aeromonas) and is diseases in fish. From the mid-1970s until nowadays, an enor- now available for collecting and sharing information about mous explosion in the number of proposed species has been Aeromonas strains from different laboratories all over the seen, and the list of species assigned to the genus is constantly world. changing. This is mainly due to two reasons: (1) the general and recent tendency to propose new species based upon single strains, especially in the last 5 years, and (2) the invalidity of Aeromonas in the Environment some species names or the use of heterotypic synonyms of previously published species. Aeromonas are described as ubiquitous bacteria. They can be To date, there are 27 valid published species names among isolated not only from a variety of aquatic environments and Aeromonas spp. included in the List of Prokaryotic names with from different terrestrial ecosystems, such as food, inverte- Standing in Nomenclature (Table 1), but the second edition of brates, plants, and slurry and fecal contents of farm animals, Bergey’s Manual of Systematic Bacteriology (Bergey’s) recognizes but also as a digestive tract symbiont of fish, leeches, and bats. far fewer. The genome sequences of 46 Aeromonas strains, Initially, three Aeromonas genomospecies (A. hydrophila, A. including both draft and complete genomes, are now available caviae, and A. veronii) were considered to be related to the vast in GenBank. majority to human infections, while A. salmonicida has been Encyclopedia of Food and Health http://dx.doi.org/10.1016/B978-0-12-384947-2.00013-1 61 62 Aeromonas Table 1 List of the valid and proposed species in the genus Aeromonas in Water Aeromonas The main reservoir of the genus Aeromonas has always been the Synonym aquatic environment, with isolates from rivers, lakes, ponds, Species Year of proposal (year of proposal) seawater (estuaries), drinking water, groundwater, wastewater, and sewage in various stages of treatments. A. hydrophila 1943 Many studies have demonstrated the ability of Aeromonas to A. salmonicida 1953 survive and grow in drinking water supplies. The bacterium can A. sobria 1981 A. media 1983 resist to water treatment strategies such as rapid/slow sand A. caviae 1984 A. punctata (1957) filtration, hyperchlorination/direct filtration, and the use of A. veronii 1988 A. ichthiosmia (1991), granular activated carbon. Studies indicated that after disinfec- À A. culicicola (2002) tion with 1 mg l 1 of chlorine, 10% of the pipes had aeromo- A. eucrenophila 1988 nads and that A. hydrophila in biofilms could survive up to À A. schubertii 1989 0.6 mg l 1 of monochloramine, which could remove E. coli A. enteropelogenes 1991 A. trota (1992) biofilms. Some studies reported that the presence of Aeromonas A. allosaccharophila 1992 in drinking water could lead to septicemia in immunocompro- A. jandaei 1992 mised persons, although no link has been demonstrated so far. A. encheleia 1995 Due to the prevalence of Aeromonas in drinking water, the A. bestiarum 1996 A. popoffii 1997 onset of new resistance mechanisms, and the presence of sev- A. simiae 2004 eral virulence factors, aeromonads are included in the ‘Con- A. molluscorum 2004 taminant Candidate List’ by the Environmental Protection A. bivalvium 2007 Agency. The World Health Organization lists Aeromonas in A. aquariorum 2008 the third edition of Guidelines for Drinking-water Quality. A. tecta 2008 On the basis of the Consumer Confidence Report Rule, public A. diversa 2010 water systems are required to report unregulated contaminants, A. fluvialis 2010 such as Aeromonas, when detected. Moreover, the presence of A. piscicola 2010 aeromonads in water supplies poses risk factors for the trans- A. sanarellii 2010 mission of these bacteria to food products such as ready-to-eat A. taiwanensis 2010 A. rivuli 2011 vegetables. Decontamination with a lactic acid solution and A. australiensis 2013 not chlorine seems to show the highest potential to reduce A. cavernicolaa 2013 Aeromonas spp. and to guarantee prolonged shelf lives of fresh-cut vegetables. aNot yet included in the species with standing in nomenclature. Aeromonas in Animals included as the predominant species in fish and water samples. Animals represent a very frequent reservoir for the transmis- However, A. hydrophila and A. veronii have been also recognized sion of Aeromonas species in the environment. Aeromonads are as involved in fish diseases, resulting in enormous economic implicated in infections of both aquatic and terrestrial organ- losses. Some studies have also identified the presence of less isms. A. salmonicida causes fish furunculosis, especially in frequently encountered species in environmental samples, salmonids, and the disease has several presentations, from an such as A. schubertii in organic vegetables. However, although acute form characterized by septicemia with hemorrhages at Aeromonas are still described as ubiquitous, the preferential the bases of fins, inappetence, and melanosis to a chronic association and adaptation between particular species and variety in older fish, consisting of lethargy, slight exopht- defined habitats have been recently highlighted. Two main halmia, and hemorrhaging in muscle and internal organs. A. different habitats were identified for Aeromonas species: aquatic hydrophila and A. veronii cause similar diseases,
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