Martn etal. D E S Drummond La Storia O V C These include L.casei, Lactobacillus paracasei, L.plantarum/paraplantarum, Lactobacillus rhamnosus and Lactobacillus curvatus, L.brevis, Lactobacillus sake, Lactobacillus pentosus, Lactobacillus acidophilus, Lactobacillus reuteri, Lactobacillus johnsonii, Lactobacillus crispatus, L.fermentum, L.buchneri and Lactobacillus gasseri. WM Hewitt H B-Z Mengesha Brinster Ruiz-Fons Bitar Mizumachi Milk and dairy products are important staples of a healthy diet. Some taxa that were consistently found across all samples included species of Streptococcus, Staphylococcus, Serratia, Pseudomonas, Corynebacterium, Ralstonia, Propionibacterium, Sphingomonas and Bradyrhizobiaceae (Fig. O (Mathieu etal., 1993; Casaus etal., 1997; Giraffa & Carminati, 1997; Achemchem etal., 2006; Izquierdo etal., 2009; Mirhosseini etal., 2010) and are active against many spoilage and pathogenic microorganisms. In Yung The possible presence of pathogenic or spoilage microorganisms in human breast milk also needs to be considered in the context that mothers may have to store milk for a variety of different reasons. H F However, recent studies have indicated that bacteria present in the maternal gut can reach the mammary gland by vertical transfer (Albesharat etal., 2011). Fitzgerald Landgren & Alvarez MA (, Lafarge Allergy to cows' milk affects 2.5% of children below 3years of age due to the presence of caseins and -lactoglobulins (Cocco etal., 2003; Gaudin etal., 2008). K McKee C RP Loneragan & Bottero MT (, Fotou etal. A recent review, which examined multiple reports of milkborne pathogen detection in bulk tanks throughout different countries, found that the percentage of tanks containing the different pathogens varied greatly. S Torriani Medina Rodriguez-Lazaro RM & Mira A (, Cailliez-Grimal Espeche K D Nahvi (, Goetsch The presence of high numbers of Gram-negative bacteria in milk has been noted in situations where hygiene standards are low and generally reflect poor udder preparation, poor sanitation or deficiencies with respect to the hygiene of equipment. Foschino A L C The latter genes in addition to those referred to above with respect to ND03 encode, for example, bacteriocins, efflux/uptake pumps, proteins involved in peptide metabolism, phage proteins as well as phage resistance genes (Rasmussen etal., 2008). Ferreira M T E J Daube Meriluoto IS Saunier I J E Beresford S P R Prevost When a model cheese system was employed to assess the consequences of the presence of some of these Gram-negative bacteria, it was established that the majority had little influence on colour, odour and volatile compounds (Delbs-Paus etal., 2011). Saffar & Nockler K (, Falentin & Poznanski E (, Franz Rossetti Moschetti Another genus detected in raw milk, Carnobacterium, consists of 11 species (www.bacterio.cict.fr; Cailliez-Grimal etal., 2005). Renckens Propionibacterium freudenreichii serves as a starter in Swiss-style cheeses. The predominant organisms in these starter cultures are lactic acid bacteria (LAB), for example, Lactococcus lactis, Lactobacillus species, Streptococcus thermophilus, Bifidobacterium species, and Leuconostoc species. & Neviani E (, Sarno MI possess a gene set involved in iron transport (Monnet etal., 2010). In laboratory-based experiments, strains of this bacterium have been shown to potentially survive pasteurisation and thus may be part of the microbial populations in both raw and pasteurised milk as well as in subsequent products (Giraffa, 2003; Ladero etal., 2011; McAuley etal., 2012). ML G In naturally fermented camel milk, Streptococcus infantarius ssp. M Lactobacillus species play a major role in fermented dairy products and also contribute to the therapeutic aspects of human health. cremoris SK11 strain contains a number of unique plasmids, pSK11A, pSK11B, pSK11L and pSK11P, which encode important traits related to dairy adaptation and utilisation, including lactose utilisation, a complex proteolytic system and an oligopeptide permease system (Siezen etal., 2005). & ul Haque I (, Gurung etal. & Alatossava T (, Murphy V These bacteria are widespread in nature, including soil and water, and are opportunistic pathogens in humans; multidrug-resistant strains are a serious concern (Dijkshoorn, 2013). Due to its long documented use in cheese manufacture, P.freudenreichii has a GRAS status (Cousin etal., 2011). Kilcawley H The absence of virulence genes from these genomes is consistent with their generally regarded as safe' (GRAS) status (Wegmann etal., 2007). & Iacono M (, Espeche (20%), Streptococcus macedonicus (10%) and Lactococcus lactis (10%; Fig. HG Although many genera of streptococci are pathogenic, S.thermophilus carries a GRAS status (Facklam, 2002) and is frequently isolated from dairy environments, including raw milk, natural starter cultures and cheese curds (Duthoit etal., 2005; Randazzo etal., 2006; Santarelli etal., 2008). As with other milk-associated microorganisms, genomic studies suggest that the genome of L.bulgaricus is in an active state of gene elimination and size reduction but does contain genes encoding complete transport systems for lactose as well as mannose, glucose, fructose and glycerol (van de Guchte etal., 2006). M Monsallier Bacteria of the genus Acinetobacter isolated from raw milk have also exhibited antibiotic resistance (Gurung etal., 2013). Tessier N Smidt Chamba Xu & Chaintreau A (, Pacini Ingestion of these microorganisms can lead to illnesses of varying severity. D Intrinsic resistance is a natural characteristic of a microorganism that allows it to grow in the presence of the corresponding antibiotic. A Nonetheless, further investigations are required to definitively establish the safety status of the species with respect to its use in dairy products. Korakli Garcia Other streptococci that have been associated with milk and milk products include Streptococcus uberis, Streptococcus agalactiae, Streptococcus dysgalactiae Streptococcus bovis and S.macedonicus. Cows' milk is produced on a massive scale. The characteristic trait of P.freudenreichii is the fermentation of lactate into propionate, acetate and CO2, while associated distinctive flavours arise from the formation of fatty acids, through lipolysis, and of branched-chain acids from the catabolism of amino acids. The microbial composition of milk can also have health-related implications in that the consumption of raw milk contaminated with pathogens can lead to, in some cases, severe illness (Oliver etal., 2009). Lactobacillus species, present in the microbiota of breast milk, is a probiotic that deserves significant attention. FJ Heilig E M Dua P However, this process also reduces the number of microorganisms that would typically contribute to desirable sensory properties associated with raw milk cheeses. & Nomura M (, Han (, Delbs-Paus & Smit G (, Yabrir Its importance in dairy products is due to its ability to rapidly convert lactose to lactate; bringing about a rapid decrease in pH; and the production of important metabolites including low levels of formate, acetoin, diacetyl, acetaldehyde and acetate (Ott etal., 2000). have the ability to produce gas (CO2), which is responsible for eye formation in some artisanal raw milk or blue-veined cheeses (Cardamone etal., 2011); metabolise lactose and citrate; and produce lactate, acetate, ethanol, acetaldehyde, diacetyl, acetoin and 2,3-butanediol, which contribute to the organoleptic properties of fermented dairy products (Vedamuthu, 1994; Sanchez etal., 2005). Although Gram-negative bacteria are regularly considered as indicators of poor hygiene and may constitute a health risk if pathogenic species are present, some may play roles in dairy fermentations by contributing positively or negatively to the sensory quality of dairy products (Delbs-Paus etal., 2011, 2012). SC M L A & Seegers H (, Gul Mavrogonatou H R While the milk, sourced from a variety of commercial producers throughout Ireland, was dominated by Lactococcus, Pseudomonas and Leuconostoc, we also detected a number of anaerobic taxa, including Bacteroides, Faecalibacterium, Prevotella and Catenibacterium, which are more typically associated with the gut microbiota and may be entering the milk through faecal contamination. T V S lactis), streptococci (such as S.salivarius) and leuconostoc (all at 102107CFUmL1; Khedid etal., 2009). R & Beyer K (, Cocolin L E Other milk and dairy isolates that exhibit probiotic properties include strains of Lactococcus lactis as well as a variety of Pediococcus, Leuconostoc, Enterococcus and Streptococcus isolates (Kim etal., 2006; Premalatha & Dhasarathan, 2011; Espeche etal., 2012; Floros etal., 2012; Forghani etal., 2012). This is the case for the Lactobacillus bacteria: they form the High prevalence of MAP has been reported in raw milk in developed countries (Argentina 8.3%, Czech Republic 2%, Ireland 0.3%, UK 6.9%, USA 028.6%; Slana etal., 2008). J Castro G Zoumpopoulou (, Afzal Litopoulou-Tzanetaki (, Zivkovic C Examples of mycotoxins of greatest public health and agro-economic significance include aflatoxin, trichothecenes, zearalenone, fumonisins and ochratoxin. JK Cai H DA Gibrat Carlier M J F Leveau F Raw milk LAB isolates also produce organic acids, hydrogen peroxide and diacetyl (Stiles, 1996). S (, Foulqui Moreno The occurrence of Salmonella ranged between 0% and 11.8% of milk samples in the USA/Canada and between 1.4% and 4% of samples in Asia, while the percentage of milk samples that was positive for the presence of Listeria ranged from 0% to 7% in USA/Canada and from 0% to 1.9% in Asia. Leuconostoc (Leuconostoc citreum, Leuconostoc lactis and Leuconostoc mesenteroides) were detected at levels of 105108CFUg1, fungal populations were present between 105 and 106CFUg1 and Enterobacteriaceae, including Enterobacter kobei, were at 103106CFUg1, but were reduced during processing. (, Deegan (, Zhang Oh Revol-Junelles These are contained within a large insertion island and encode a transposase, a glutamate decarboxylase, an acetyltransferase, a polysaccharide biosynthesis protein and proteins associated with exopolysaccharide biosynthesis (Sun etal., 2011). & Topisirovic L (, Olivares Guss MA O T Rizzotti GN & Wilkinson MG (, Hunt Caron E D & Tzanetakis N (, Walther Of those which could be identified, many corresponded to technologically important bacteria such as Lactobacillus, Leuconostoc and Enterococcus spp. CL MD BG Terzic-Vidojevic Popoff MC 2). Atun Zanetti L After a century of nisin research, where are we now and where are we going? Moraes K F L & Garca-Lpez MaL (, Arora It is important to understand the changes that can occur in the microbiology of raw milk during its storage and as a consequence of subsequent treatments. Mycotoxins, that is, low molecular weight compounds produced as secondary metabolites by filamentous fungi, can lead to illness in humans with symptoms such as nausea, vomiting, diarrhoea and headache (Creppy, 2002), and some mycotoxins have carcinogenic potential (Murphy etal., 2006). A C Holzapfel It is defined as homofermentive lactic acid bacteria Thus, lactobacilli numerically overshadow all other genera, including Bifidobacterium, in colonizing neonatal mucosal and skin surfaces at birth during passage though the vagina and its Lactobacillus -rich microbiota, and then after labour during feeding, with the mothers breast milk containing high numbers of lactobacilli [ 5 ]. Mukerji I Gatti C L These might also be regarded as biopreservative agents that could be used to extend the shelf life and safety of fermented and other foods (Stiles, 1996). Zeng WebLactobacillus bulgaricus is one of over 200 published species in the Lactobacillus genome complex (LGC) [not verified in body] and is the main bacterium used for the production of yogurt.It also plays a crucial role in the ripening of some cheeses, as well as in other processes involving naturally fermented products. Strains of P.freudenreichii, and to a lesser extent P.acidipropionici, have begun to attract attention as potential probiotics as a consequence of studies revealing an ability, either alone or in combination with other probiotics, to reduce pathogen adhesion to mucus (Collado etal., 2008), increase bifidobacteria counts in the gut, aid in restoring a healthy gut microbiota, improve bowel movement, alleviate inflammatory disorders and reduce allergy development in infants (Jan etal., 2002; Cousin etal., 2012). lactis (Giraffa etal., 2008). Some potential raw milk-derived bacteriocins have been characterised in-depth. have the ability to survive on surfaces, tools and pasteurisers for long periods of time and to resist heat treatments and refrigeration temperatures (Hemme & Foucaud-Scheunemann, 2004). etal. Donkey milk is less commonly consumed and, for the same reason, is less extensively studied. Carnobacteria are slow acidifiers and therefore are not suitable for the use as starter cultures but can be considered to be beneficial NSLAB due to their aromatic and flavour contributing end products (Afzal etal., 2010). More recently, through the application of high-throughput sequencing, an even more diverse population has been uncovered (Table4; Hunt etal., 2011; Cabrera-Rubio etal., 2012). & Jakobsen M (, Mathieu T JM Schoenfeld (, Liatsos NGS results are represented as a percentage of total reads. A & Sorokin A (, Bonizzi protection against diarrhoea), immunostimulatory and anticarcinogenic capabilities, lowering of serum cholesterol and alleviation of lactose maldigestion (Fernndez etal., 2012). Rodriguez There are more than 200 strains of bacteria in breast milk, of which the most important are Lactobacilli, Bacteroides and Bifidobacterium. Genome sequencing of Lactococcus lactis ssp. Capita Causation or simple association? Kluyveromycesmarxianus is of particular interest due to its fast growth rate, thermotolerance, the ability to assimilate a wide range of sugars, the secretion of lytic enzymes and the production of ethanol by fermentation (Lane & Morrissey, 2010). Duthoit A JA Belzacq In a study investigating the presence of Gram-negative bacteria in different cheeses produced in France, 173 isolates were isolated. WebLactobacillus Lactobacillus SPP. G Another, relatively new, pathogen of concern to the dairy industry is Shiga-toxin-producing E.coli (STEC). Markman While these microorganisms are naturally present in raw milk and artisanally produced cheeses (Gaya etal., 2001), they are frequently added to pasteurised milk to facilitate the commercial manufacture of cheeses (Smit etal., 2005). These populations, which are often referred to as nonstarter LAB (or NSLAB), have an ability to adapt to the conditions of cheese ripening, where many nutrients are depleted, pH is reduced, and moisture content is low. The mould genera that are most commonly detected in raw milk include Penicillium, Geotrichum, Aspergillus, Mucor and Fusarium (Lavoie etal., 2012). ML Pot A Psychrotrophic populations, which These viruses infect bacteria and, after intracellular replication, lyse their host cells (Marc etal., 2012). Santoro P While L.bulgaricus encodes a full set of genes for biosynthesis of folate, it is thought that S.thermophilus is required to produce p-aminobenzoate, which feeds this pathway. etal. TP MG Notably, some of the latter would be typically regarded as thermophilic microorganisms. & Lindqvist R (, Ruaro F Microorganisms can also negatively impact on milk quality and shelf life; for example, psychrotolerant bacteria can proliferate during refrigeration and, through the production of extracellular lipases and proteases, result in spoilage (Desmasures & Gueguen, 1997; Hantsis-Zacharov & Halpern, 2007). P AK Pepe V Mauriello There is evidence to suggest that raw milk purchased from retailers represents a greater risk of Listeria-associated illness than milk obtained directly from milk tanks on farms, most likely as a consequence of the growth of the pathogen over the extended storage period (Latorre etal., 2011). & Hernandez M (, Madera lactis IL1403 revealed that all known genes required for energy metabolism were present, including a number of genes involved in fermentation as well as a novel gene, poxL, encoding pyruvate oxidase, which may play a role in switching between fermentation modes. cremoris in particular, are primarily known for their role as starter cultures for the cheese industry. ML Ricci C Scarpellini Martinez Comunian These microorganisms are distinguished from one another on the basis of arginine and citrate utilisation, growth temperature and salt tolerance (Kahala etal., 2008). I Daga Overall, 26 different genera were present. More recently, high-throughput sequencing has been applied to identify the bacterial populations present in buffalo milk and throughout the manufacture of mozzarella cheese (Table3; Ercolini etal., 2012). M Moschetti M Barbour Siezen Nearly half of all isolates were representatives of Enterobacteriaceae. Silva Jnior Lauret K Mutharia Wang J A J Traditionally, S.aureus has been considered as the main causative agent; however, Staphylococcus epidermidis is emerging as a leading cause of subacute and acute mastitis in both women and veterinary medicine (Delgado etal., 2009). MH A Fadda & Muzquiz JL (, Verdier-Metz lactis phenotype and 10 of the ssp. RJ J Ma The growth of L.helveticus in milk is dependent on a complex system of proteolytic enzymes, which collectively enable strains to access essential amino acids (Christensen etal., 1999). Bobo Glass & Cotter PD (, Raats JP The second region corresponds to a prophage. Di Palo E Lactococcus, Lactobacillus, Streptococcus, Propionibacterium and fungal populations), causing spoilage (e.g. & Von Mutius E (, Bravo RR AM RJ Rodriguez-Benitez SAY & Engels WJM (, Sun Zhao W Franzetti Ramos Wilkinson Milk is a highly nutritious food that can be obtained from a variety of animal sources such as cows, goats, sheep and buffalo, as well as humans, for human consumption. JR Civera Hatzikamari M P D Indeed, L.helveticus is currently used in the production of fermented drinks such as Evolus (Valio Ltd., Valio, Finland) and Calpis (Calpis Food Industry Co. Ltd., Tokyo, Japan), which have properties associated with a reduction in blood pressure through the inhibition of ACE as a consequence of the production of bioactive tripeptides (Slattery etal., 2010). & Pisano MB (, Coton The antihypertensive properties of these microorganisms are being investigated and exploited by industry with a view to producing health-promoting drinks. Andrighetto The first contains genes involved in cell wall biosynthesis. Sheep milk is rarely consumed but still constitutes c.1.3% of global milk production as it is often employed throughout Europe in the development of cheese (Tsakalidou & Odos, 2012). One must also consider the use of strategies, such as the use of DNA-binding agents or an alternative focus on RNA, to limit the risk of false positives resulting from the amplification of DNA from dead cells (Quigley etal., 2011). J etal. Sandoval H during outdoor feeding (Hagi etal., 2010), on the location of the animals (Bonizzi etal., 2009) and on the lactation stage (Callon etal., 2007). Finally, it is notable that the implementation of strict hygiene standards brings about a reduction in the microbial load of milk, including a reduction in populations of technological importance, which can, in turn, impact negatively on cheese manufactured using traditional or artisanal approaches (Monsallier etal., 2012). L I There was also a large percentage of unassigned reads (c.20%) corresponding to the raw milk. Zeng WJ & Fortina MG (, Florez Macioniene Buffoni It furthers the University's objective of excellence in research, scholarship, and education by publishing worldwide, This PDF is available to Subscribers Only. E The most recent classification groups LAB bacteriocins into two classes; class I are post-translationally modified bacteriocins, and class II are unmodified bacteriocins (Rea etal., 2011). However, the fact that one of the studies referred to above revealed that c.50% of the Gram-negative strains isolated were resistant to several antibiotics is a particular cause for concern (Delbs-Paus etal., 2011). Bacterial populations detected in raw milk of sheep or buffalo using culture-dependent, culture-independent and next-generation DNA sequencing methods, Bacterial populations detected in raw human milk using culture-dependent, culture-independent and next-generation DNA sequencing methods. Van Landschoot M Peptidases and amino acid catabolism in lactic acid bacteria, Mutational analysis of major, sequential IgE-binding epitopes in alpha(s1)-casein, a major cow's milk allergen, The late blowing in cheese: a new molecular approach based on PCR and DGGE to study the microbial ecology of the alteration process, Risk factors for pulmonary tuberculosis in Russia: casecontrol study, Adhesion and aggregation properties of probiotic and pathogen strains, Comparative analysis of the diversity of aerobic spore-forming bacteria in raw milk from organic and conventional dairy farms, Antilisterial activity of nisin-like bacteriocin-producing, Diversity and assessment of potential risk factors of Gram-negative isolates associated with French cheeses, Bacteriocins: developing innate immunity for food, Dairy propionibacteria as human probiotics: a review of recent evidence, Update of survey, regulation and toxic effects of mycotoxins in Europe, Origins, acquisition and dissemination of antibiotic resistance determinants, Short communication: influence of storage and preservation on microbiological quality of silo ovine milk, Influence of storage conditions on the growth of, Bacteriocins: biological tools for bio-preservation and shelf-life extension, Fungal diversity in cow, goat and ewe milk, Monitoring bacterial communities in raw milk and cheese by culture-dependent and -independent 16S rRNA gene-based analyses, Benefits and risks associated with Gram-negative bacteria within cheese microbial communities, 10th International Meeting on Mountain Cheese 1415 September 2011, Ed. M It is generally accepted that the lactic acid bacteria (LAB), a group of bacteria that ferment lactose to lactate, are a dominant population in bovine, goat, sheep and buffalo milk, prior to pasteurisation. Papandreou I R & Malik RK (, Binetti Capra C
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