A Review of Salmonella Enterica with Particular Focus
Total Page:16
File Type:pdf, Size:1020Kb
Veterinary World, EISSN: 2231-0916 REVIEW ARTICLE Available at www.veterinaryworld.org/Vol.12/April-2019/5.pdf Open Access A review of Salmonella enterica with particular focus on the pathogenicity and virulence factors, host specificity and antimicrobial resistance including multidrug resistance Saleh Mohammed Jajere Department of Veterinary Public Health and Preventive Medicine, Faculty of Veterinary Medicine, University of Maiduguri, PMB 1069, Maiduguri, Borno State, Nigeria. Corresponding author: Saleh Mohammed Jajere, e-mail: [email protected] Received: 19-12-2018, Accepted: 11-02-2019, Published online: 06-04-2019 doi: 10.14202/vetworld.2019.504-521 How to cite this article: Jajere SM (2019) A review of Salmonella enterica with particular focus on the pathogenicity and virulence factors, host specificity and antimicrobial resistance including multidrug resistance, Veterinary World, 12(4):504-521. Abstract Salmonella genus represents the most common foodborne pathogens frequently isolated from food-producing animals that is responsible for zoonotic infections in humans and animal species including birds. Thus, Salmonella infections represent a major concern to public health, animals, and food industry worldwide. Salmonella enterica represents the most pathogenic specie and includes > 2600 serovars characterized thus far. Salmonella can be transmitted to humans along the farm-to-fork continuum, commonly through contaminated foods of animal origin, namely poultry and poultry-related products (eggs), pork, fish etc. Some Salmonella serovars are restricted to one specific host commonly referred to as “host-restricted” whereas others have broad host spectrum known as “host-adapted” serovars. For Salmonella to colonize its hosts through invading, attaching, and bypassing the host’s intestinal defense mechanisms such as the gastric acid, many virulence markers and determinants have been demonstrated to play crucial role in its pathogenesis; and these factors included flagella, capsule, plasmids, adhesion systems, and type 3 secretion systems encoded on the Salmonella pathogenicity island (SPI)-1 and SPI- 2, and other SPIs. The epidemiologically important non-typhoidal Salmonella (NTS) serovars linked with a high burden of foodborne Salmonella outbreaks in humans worldwide included Typhimurium, Enteritidis, Heidelberg, and Newport. The increased number of NTS cases reported through surveillance in recent years from the United States, Europe and low- and middle-income countries of the world suggested that the control programs targeted at reducing the contamination of food animals along the food chain have largely not been successful. Furthermore, the emergence of several clones of Salmonella resistant to multiple antimicrobials worldwide underscores a significant food safety hazard. In this review, we discussed on the historical background, nomenclature and taxonomy, morphological features, physical and biochemical characteristics of NTS with a particular focus on the pathogenicity and virulence factors, host specificity, transmission, and antimicrobial resistance including multidrug resistance and its surveillance. Keywords: Enteritidis, foodborne pathogens, Heidelberg, multidrug-resistant, pathogenicity and virulence factors, Salmonella enterica, Typhimurium. Introduction non-motile, the majority of members in the genus Salmonella represents a large genus of global Salmonella are motile by peritrichous flagella. The public health significance and is the leading cause SG and SP are associated with clinical disease in of foodborne illnesses responsible for thousands poultry, and they cause considerable economic losses of deaths worldwide [1-9]. Salmonella is Gram- – due to the replacement of infected flocks and asso- negative, rod-shaped bacteria, and facultative anaer- ciated treatment costs – to poultry farmers, especially obes belonging to the family Enterobacteriaceae. in developing countries of the world [11-13]. In gen- The genus Salmonella belongs to two broad species eral, the genus has a predilection limited to the diges- namely Salmonella enterica and Salmonella bon- tive tracts of both humans and animals hosts. Thus, gori. So far, more than 2600 serovars belonging to S. the presence of Salmonella in other habitats such as enterica have been described worldwide, and many the water, environment, and food represents fecal of these serovars are capable of causing illnesses in contamination. Recent data from the United States, both humans and animals [10]. While few variants of European countries, and low- and middle-income S. enterica namely Salmonella Gallinarum (SG) and countries (LMICs) indicate that Salmonella cases are Salmonella Pullorum (SP) are non-flagellated and the most commonly encountered cause of bacterial foodborne disease globally and hence supporting the Copyright: Jajere. Open Access. This article is distributed under fact that the control programs aimed at reducing the the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which Salmonella contamination along the food chain have permits unrestricted use, distribution, and reproduction in any not been successful [14]. Consequently, there will be medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons increased frequency and persistence of S. enterica in license, and indicate if changes were made. The Creative Commons the intestinal tracts of food animals and this situation Public Domain Dedication waiver (http://creativecommons.org/ publicdomain/zero/1.0/) applies to the data made available in this creates chronic or non-symptomatic carriers that con- article, unless otherwise stated. tinue to shed the organism in their feces. Thus, these Veterinary World, EISSN: 2231-0916 504 Available at www.veterinaryworld.org/Vol.12/April-2019/5.pdf carriers serve as reservoirs for future contamination the organism and subsequently Gaffky isolated the and spread of Salmonella by contaminated milk, meat, bacillus responsible for human typhoid fever [55]. eggs and other agricultural products fertilized and Thereafter, in 1885 Theobald Smith together with grown in Salmonella contaminated manure [14]. Daniel Elmer Salmon discovered and isolated In recent years, the development of antimicrobial Salmonella from the intestines of pigs infected with resistance (AMR) among foodborne pathogens such classical swine fever (hog cholera). During this period, as Salmonella have been associated with an increased they thought the bacterium was the etiological agent number of human deaths, longer duration of hospi- of hog cholera [55,56]. Later, the bacterial strain was tal stay, and high costs of treatment due to therapy named Salmonella after Dr. Daniel Elmer Salmon, an failure. Several clones of multidrug-resistant (MDR) American pathologist who has worked together with Salmonella have emerged during the late 1990s and Smith [57]. In recent years, the issue of nomencla- early 2000s and since then, their prevalence both in ture of the genus Salmonella has been complex, con- humans, domestic animals and other wildlife spe- troversial, and still remains subject of debate [57]. cies have expanded globally [9,15-25]. Recently, the At present, most Salmonella reference centers in increasing prevalence of MDR Salmonella such as the world including the Centers for Disease Control resistance towards clinically important antimicrobials (CDC) adopt the nomenclatural system of Salmonella like fluoroquinolones and third-generation cepha- as recommended by the World Health Organization losporins has become an emerging problem world- (WHO) [58]. This nomenclatural system classifies the wide [16,18,26-33]. In recent years, studies on the genus Salmonella into two species based on differ- global burden of non-typhoidal Salmonella (NTS) ences in their 16S rRNA sequence analysis. These two have shown an increasing incidence of NTS. For broad species included S. enterica (type species) and instance, one of these studies estimated that there are S. bongori [57]. approximately 94 million cases of NTS gastroenteritis Nomenclature/Taxonomy resulting in 155,000 deaths globally each year [34]. According to this study, majority of the NTS burden On the basis of biochemical properties and was found in the Southeast Asian and the Western genomic relatedness, the S. enterica is further clas- Pacific regions [34,35]. sified into six subspecies [59]. These subspecies in Moreover, of the 94 million NTS cases the nomenclature are denoted by Roman numerals: reported [34], 80.3 million cases were estimated as I. S. enterica subsp. enterica; II. S. enterica subsp. foodborne origin [15]. Among the NTS, Salmonella salamae; III. S. enterica subsp. arizonae; IIIa. S. enter- Typhimurium (ST), Salmonella Enteritidis (SE), ica subsp. diarizonae; IV. S. enterica subsp. houtenae; Salmonella Heidelberg (SH), and Salmonella Newport and V. S. enterica subsp. indica. Of all the subspecies (SN) are the epidemiologically important NTS sero- of Salmonella, the S. enterica subsp. enterica (I) is types – with poultry and poultry derived products as the most common and is found predominantly asso- important reservoir sources – and have been associ- ciated with mammals and attributes about 99% of ated with the majority of human salmonellosis burden Salmonella infections in humans and warm-blooded worldwide [5,36-40]. S. enterica is widely distributed animals. On the other hand, the other five