Staphylococcus aureus

JA Hudson, Food Safety Programme, ESR, Christchurch, New Zealand

r 2014 Elsevier Ltd. All rights reserved.
This article is a revision of the previous edition article by JA Hudson, volume 2,
pp 820–825, © 2004, Elsevier Ltd.

Glossary Fail-safe Used in predictive microbiology to represent a

aw (water activity) The available water in a given situation where a model predicts growth but none occurs.
substrate. Values range from 0 to 1, where 1 is aw of pure The model has failed but in a safe manner.
distilled water. Humectant An additive that binds water and controls
D-value The decimal reduction time, or the duration of an water activity.
antibacterial process (such as heat or irradiation) at a given Microflora The naturally occurring microorganisms in a
intensity which results in a 90% reduction in the bacterial given environment.
population.

Introduction For example, limited growth of S. aureus occurred on vacuum-

packed ham and turkey, but not on chicken stored at 10 °C.
The first link between Staphylococcus aureus and food poisoning The optimum pH for growth is 7.0–7.5 with a range of 4.2–
was made following an outbreak associated with eating cheese, 9.3. The lower pH value at which growth occurs is raised when
and the first recognized meat-related outbreak, involving a organic acids such as acetic and lactic acids are used as the
fatality, was reported in 1894. The organism has subsequently acidulant, and no growth occurs at room temperature in vac-
been incriminated in incidents involving a wide range of food uum-packaged fermented meat products of pH ≤ 5.3. Growth
vehicles including meat, poultry, dairy, cream-filled bakery, is optimal under aerobic conditions but the organism can also
and egg products as well as salads and canned mushrooms. grow anaerobically. Of note is the organism's ability to grow at
Staphylococcal food poisoning is an intoxication caused by low aw values. Growth can occur at aw values as low as 0.86
the consumption of one of a variety of enterotoxins. A number (equivalent to 20% NaCl) depending on the humectant used
of species of Staphylococcus are able to produce these toxins but (e.g., NaCl, sucrose, etc.), and the organism grows well in the
only one species, S. aureus, is commonly associated with presence of 7–10% NaCl.
foodborne disease. It is regarded as an organism that competes Staphylococcus aureus is moderately heat resistant for a
poorly with the natural microflora of raw meats but is able to nonspore-forming foodborne pathogen, with a mean D-value
grow at low water activity (aw) values. These two characteristics (i.e., time for a 90% reduction) of approximately 5–7 min at
are reflected in the types of meat products normally associated 60 °C. However, in salty foods its thermal resistance is much
with staphylococcal foodborne disease: i.e., foods that have greater. For example, the mean D-value at 60 °C increased
been cooked and hence have no competitors present; and low from 6 to 25 min when the NaCl content of meat macerate
aw foods, often with high salt concentrations, which are con- was increased from 0% (w/v) to 8.4%. Microwave heating to
ducive for the growth of S. aureus. 65 °C results in reductions of numbers of the order of 1.7–
The toxin produced is heat stable and can survive com- 2.5 log10 cfu ml−1 or g−1 in a variety of foods. D-values for
mercial canning. Staphylococcus aureus may, therefore, be ab- irradiation are in the range of 0.3–0.6 kGy depending on the
sent from a food, yet the food can still contain the toxin and atmosphere, temperature and nature of the food substrate in
cause disease. Good food handling practices (e.g., good per- which the organism is irradiated. However, D-values as high as
sonal hygiene, prevention of cross-contamination and tem- 0.86 kGy have been reported.
perature control) are the most effective way of preventing For a nonspore-forming organism, S. aureus is very resistant
staphylococcal intoxications. to adverse conditions and it can survive long periods of des-
iccation. For example, the organism can survive for more than
1000 days on a dry plastic surface. Survival during frozen
storage is good at temperatures ≤10 °C, although there can be
Characteristics of the Organism and its Toxin
some loss of viability during freezing.
Staphylococcus aureus is frequently found in meat products,
The Organism
but usually at low concentrations. Some examples are given in
Staphylococcus aureus is a Gram-positive, catalase-positive coc- Tables 1 and 2. Data are presented for detection of the or-
cus, 0.5–1.5 mm in diameter, which forms clusters of cells ganism and toxin by conventional methods, but reported
appearing as characteristic ‘bunches of grapes’ when viewed prevalence rates are higher when molecular methods such as
microscopically. In laboratory media it has an optimum the polymerase chain reaction (PCR) are used.
temperature for growth of 37 °C with a range of 7–48 °C. An emerging concern with S. aureus is the acquisition of
However, when growing in food these limits may be reduced. methicillin resistance and the roles that food production and

376 Encyclopedia of Meat Sciences, Volume 2 doi:10.1016/B978-0-12-384731-7.00041-6

 Microbiological Safety of Meat | Staphylococcus aureus 377

Table 1 Examples of the prevalence of Staphylococcus aureusa on or in meat products, and the
prevalence of enterotoxin-positive strains among isolates from such products

Product Proportion samples +ve (%) Isolates enterotoxin +ve (%)

Raw pork, France 57.7 34.6

Retail smoked ham, France 11.1 0
Fresh meat, Italy 26.1 21.4
Minced meat/burgers, Italy 31.2 46
Fresh meat preparations, Italy 10.6 53.7
Fermented sausages, USA ND 0
Fish products, India 21 41
Ready-to-eat meat, Korea 2.1 100
Raw fish, Korea 19.8 50
Deboned beef, South Africa 15.8–24.4 ND
Beef, USA 20.5 ND
Chicken, USA 25.0 ND
Turkey, USA 24.6 ND
Turkey, USA 77 ND
Pork, USA 42 ND
Chicken, USA 41 ND
Beef, USA 37 ND
Boneless beef trim, USA 4.2 ND
Boneless beef trim, Australia 4.0 ND
Boneless beef trim, New Zealand 8.2 ND
Boneless beef trim, Uruguay 29.5 ND
a
Or coagulase-positive staphylococci.
Abbrevation: ND, Not determined.
Source: Atanassova, V., Meindl, A., Ring, C., 2001. Prevalence of Staphylococcus aureus and staphylococcal enterotoxins in raw
pork and uncooked smoked ham-a comparison of classical culturing detection and RFLP-PCR. International Journal of Food
Microbiology 68, 105−113; Bhargava, K., Wang, X., Donabedian, S., et al., 2011. Methicillin-reistant Staphylococcus aureus in
retail meat, Detroit, Michigan, USA. Emerging Infectious Diseases 17, 1135−1137; Bosilevac, J.M., Guerni, M.N., Brichta-
Harhay, D.M., Arthur, T.M., Koohmarie, M., 2007. Microbiological characterization of imported and domestic boneless beef trim
used for ground beef. Journal of Food Protection 70, 440−449; Levine, P., Rose, B., Green, S., Ransom, G., Hill, W., 2001.
Pathogen testing of ready-to-eat meat and poultry products collected at federally inspected establishments in the United States,
1990 to 1999. Journal of Food Protection 64, 1188−1193; Normanno, G., Firinu, A., Virgilio, S., et al., 2005. Coagulase-positive
staphylococci and Staphylococcus aureus in food products marketed in Italy. Inernational Journal of Food Microbiology 98,
73−79; Oh, S.K., Lee, N., Cho, Y.S., et al., 2007. Occurrence of toxigenic Staphylococcus aureus in ready-to-eat food in Korea.
Journal of Food Protection 70, 1153−1158; Shale, K., Lues, J.F.R., Venter, P., Buys, E.M., 2005. The distribution of
Staphylococcus sp. On bovine meat from abattoir deboning rooms. Food Microbiology 22, 433−438; Simon, S.S., Sanjeev, S.,
2005. Prevalence of enterotoxigenic Staphylococcus aureus in fishery products and fish processing factory workers. Food Control
18, 1565−1568; and Waters, A.E., Contente-Cuomo, T., Buchhagen, J., et al., 2011. Multidrug resistant Staphylococcus aureus
in US meat and poultry. Clinical Infectious Diseases 52, 1−4.

Table 2 Examples of data for concentrations of Staphylococcus consumption may have in infections of people with methi-
aureus in cooked sliced meats (A), ready-to-eat pies and pastries (B) cillin-resistant S. aureus (MRSA). Testing of Canadian feedlot
and other cooked meat products (C) cattle just before slaughter failed to detect MRSA; but MRSA
has been detected in other food animals such as pigs, and in a
Count (g−1) Fraction of samples (%) positive
variety of retail meat products. Examples are given in Table 3.
A B C Widespread or emerging resistance to other antibiotics is also a
cause for concern.
ND 91.2 96.6 100
20≤102 7.4 3.0 0
102≤103 0.9 0.3 0 Staphylococcal Enterotoxins
103≤104 0.4 0 0
104≤105 0 0.2 0 The organism causes disease through the production of 1 of a
105≤106 0.1 0 0 group of at least 20 structurally similar protein toxins. Only
106≤107 0.1 0 0 five subclasses of these toxins are commonly involved in
ND o20 CFU g−1. human disease, i.e., staphylococcal enterotoxins (SE) types A,
Source: Little, C.L., de Louvois, J., 1998. The microbiological examination of butchery B, C, D, and E, with type C toxins being further divided into
products and butchers' premises in the United Kingdom. Journal of Applied types C1, C2, and C3. Human disease usually involves SE-A.
Microbiology 85, 177−186. Toxin production occurs under more restrictive conditions
378 Microbiological Safety of Meat | Staphylococcus aureus

Table 3 Examples of data for the prevalence of methicillin-resistant S. aureus (MRSA) in meat

Country Samples (number tested) Number +ve for MRSA/Number tested Percentage positive
a
The Netherlands Beef 42/395 10.6
Veal 39/257 15.2
Pork 33/309 10.7
Lamb/mutton 20/324 6.2
Chicken 83/520 16.0
Turkey 41/116 35.3
Fowl 4/118 3.4
Game 4/178 2.2
USA Beef 2/156 1.3
Chicken 3/76 3.9
Turkey 1/57 1.7
USA Minced pork 1/300 0.3
Minced beef 0/198 0
Minced turkey 0/196 0
Germanya Fresh chicken 6/24 25.0
Chicken meat products 4/19 21.1
Fresh turkey 11/22 50.0
Turkey meat products 11/21 52.4
a
These studies used an enrichment and so were more sensitive than others using direct plating.
Source: Bhargava, K., Wang, X., Donabedian, S., et al., 2011. Methicillin-reistant Staphylococcus aureus in retail meat, Detroit, Michigan, USA. Emerging Infectious Diseases 17,
1135−1137; de Boer, E., Zwartkruis-Nahuis, J.T.M., Wit, B., Huijsdens, X.W., et al., 2009. Prevalence of methicillin-resistant Staphylococcus aureus in meat. International Journal of
Food Microbiology 134, 52−56; Feßler, A.T., Kadlec, K., Hassel, M., et al., 2011. Characterization of methicillin-resistant Staphylococcus aureus isolates from food and food products
of poultry origin in Germany. Applied Environmental Microbiology 77, 7151−7157; and Kelman, A., Soong, Y.-A., Dupuy, N., et al., 2011. Antimicrobial susceptibility of
Staphylococcus aureus from retail ground meats. Journal of Food Protection 74, 1625−1629.

than those that allow growth. It occurs between 10 and 45 °C

and at pH 4.8–9.0, and is optimal at temperatures between 35
and 40 °C and pH values between 5.3 and 7.0. Toxin pro-
duction is greater under aerobic than under anaerobic con-
ditions, and occurs at aw values ≥0.90. The toxin is very
resistant to heat. For example, the D-value for SE-B is 100 min
at 149 °C and an aw of 0.99, and 225 min when the aw is 0.90.
The toxin is also resistant to the proteolytic enzymes that occur
in the gastrointestinal tract, dehydration, gamma irradiation,
and extremes of pH.
Because the toxin is heat stable, it is possible for a food to
be free of viable organisms yet still be the cause of SE poi-
soning. This may occur if the required population was reached,
enterotoxin was produced, and the cells were then inactivated
by a process such as cooking.
Ingestion of less than 1.0 mg of toxin can result in illness,
but this level is reached only when the population of S. aureus Figure 1 Typical colonies of S. aureus growing on BP agar.
exceeds 105 g−1. Concentrations as high as 1.5×1010 cfu g−1
have been reported in outbreak-related foods and ingested
levels of 1–5 mg of toxin are normally associated with out- of the organism in a sample is needed rather than a sensitive
breaks, although children have become sick at a dose of 17 ng. detection method.
Not all isolates of S. aureus produce toxin. For example, one There is a good overall agreement with regard to the
study of coagulase-positive food isolates found that only methods that are appropriate for S. aureus (see ISO 6888-1
30.5% were unequivocally enterotoxin positive. and ISO 6888-2). Baird-Parker (BP) agar is used as a plating
medium in the ISO and many other methods. Typical
colonies on BP agar appear 2–3 mm in diameter, black and
shiny, surrounded by a white edge and a clear zone (Figure 1).
Isolation and Identification Other media such as Rabbit Plasma Fibrinogen Agar
(RPFA) are available. In this case, the medium identifies
Enumeration and Detection coagulase- and thermonuclease-positive colonies. With
Because relatively high numbers of the organism are required these media, the sample is prepared and dilutions either
to cause illness, in most cases an estimate of the concentration spread onto them (BP) or added to pour plates (RPFA),
 Microbiological Safety of Meat | Staphylococcus aureus 379

the plates are incubated and suspect colonies counted and manufacturers. In addition, kits are available for the detection
confirmed. A Most Probable Number method using enrich- of the SE. While being too numerous to detail, a list of ap-
ments of sample dilutions in Giolitti and Cantoni broth fol- proved test kits, along with their status in terms of recognition
lowed by plating to BP agar is recommended by the American is maintained by AOAC International.
Public Health Association (APHA). A number of other pro-
prietary media are available in conventional agar and film
formats. Typing
For detection of S. aureus in foods in which cells may be
Numerous methods have been developed for the typing of S.
injured, for example, in processed foods, the APHA recom-
aureus. Some of the methods that have been applied to meat
mend a short enrichment in a nonselective medium followed
products or outbreak investigations are briefly mentioned
by a period of incubation with NaCl present as a selective
here.
agent. Isolates are then obtained on BP agar.
Isolates may be assigned to different biotypes/ecovars ac-
Several PCR methods that allow detection of S. aureus
cording to staphylokinase, β-hemolysin, coagulase, and crystal
have now been published and commercial systems based
violet agar growth tests. These ecovars are largely host-specific
on PCR are also available. Because the methods target diffe-
being grouped into human, poultry-like, bovine, ovine, and
rent genes, they vary in their suitability for various tasks.
nonhost-specific ecovars. Molecular typing produces data that
General methods target genes such as nuc which encodes
also tend to reflect a clonal association of S. aureus types and
thermostable nuclease production. Others more specifically
animal hosts. A few tests may therefore be useful in identifying
target toxin or antibiotic resistance genes. PCR methods are
the animal origin of S. aureus isolates, although a significant
available as real-time applications for the detection of the or-
proportion of isolates may be of unspecified origin.
ganism in foods, and such techniques offer the possibility of
Phage typing is useful for the differentiation of isolates, and
quantifying rather than just detecting the presence of S. aureus.
most enterotoxin-producing isolates have been shown to be-
long to a restricted range of phage types. However, the level of
expertise required to use the methodology is likely to preclude
Identification small or routine testing laboratories from using it. Coagulase
Staphylococci are usually distinguished from other Gram- typing has also been used.
positive facultatively anaerobic cocci by the presence of cata- Pulsed field gel electrophoresis (PFGE) is a highly dis-
lase in Staphylococcus. Distinction between S. aureus and the criminatory molecular typing technique that has been suc-
micrococci is more difficult but can be achieved using the tests cessfully used to type S. aureus isolates. The primary restriction
shown in Table 4. Staphylococcus aureus is characteristically enzyme used was SmaI, with KspI being used subsequently to
positive for the following tests; thermostable nuclease, coa- further evaluate indistinguishable isolates. SmaI digests pro-
gulase, clumping factor, yellow pigment, acetoin production, duced profiles with 10–20 fragments in the 20–700 kb range.
and hemolysis. The presence in S. aureus of clumping factor Clusters produced by PFGE correspond to those produced by
can be used to distinguish it from other coagulase-positive biotyping, but not phage typing. PFGE has been shown to be
Staphylococcus species. more discriminatory than amplified fragment length poly-
Not all S. aureus isolates are capable of producing toxin and morphism analysis. Staphylococcal protein A (spa) typing and
it may be necessary to demonstrate the toxin-forming cap- multilocus sequence typing are other techniques that have
ability of food isolates. Because S. aureus may have grown, been applied.
produced toxin, and then have been inactivated by cooking, a
ready-to-eat food containing low numbers of the organism
may not be safe. Toxin testing is therefore necessary in outbreak Characteristics of Foodborne Illness
investigations where there is potential for enterotoxin to have
been produced in the food before a bactericidal treatment. Symptoms are caused by the production of a toxin which is
Rapid test kits for the detection, identification, and con- thought to act by causing a signal to be sent from intestinal tract
firmation of S. aureus are readily available from diagnostics receptors to the medullary emetic center of the brain. The precise
area in the abdomen that is stimulated has not been identified.
Table 4 Biochemical tests that discriminate between S. aureus and Symptoms normally occur 0.5–6 h (usually 2–4 h) after
Micrococci ingestion of the toxin and are most often nausea, frequent
vomiting, and abdominal cramps for up to 48 h. Diarrhea is
Test S. aureus Micrococci
less common but can occur in a significant proportion of cases,
Lysostaphin resistance S R and always occurs in conjunction with vomiting. In severe
Coagulase production + − cases, headaches, fever, and collapse may occur. Recovery is
Lysozyme resistance R R/S usually swift, occurring over a few hours to 1 day, and gen-
Thermostable nuclease production + − erally no treatment is required. Death is not a common con-
Anaerobic glucose fermentation + − sequence of staphylococcal intoxication, but has been reported
Anaerobic mannitol fermentation + − in the young and elderly. The organism may also be associated
Modified oxidase − +
with autoimmune disorders. Estimates from the USA for ill-
Erythromycin resistance (0.4 mg ml−1) R S
ness associated with this organism are a hospitalization rate of
Abbreviations: R, resistant; S, sensitive. 6.4% and a case fatality rate of o0.1%.
380 Microbiological Safety of Meat | Staphylococcus aureus

Outbreaks of illness are often caused by contamination of example, pH and aw may be used in combination at levels
foods by food handlers with uncovered infected wounds. It which alone would not prevent growth of the organism. Pre-
may also be transferred to food from the nose, where it is a servatives such as potassium sorbate and potassium nitrite
commensal in 30% of the population, or other moist parts of may also be used to inhibit growth of the organism, especially
the body, for example, mucous membranes and the skin. when used in combination with another controlling factor. As
Enterotoxin production seems to be linked with isolates of an example, the inactivation rate in the presence of 2.5% po-
human origin, further strengthening the link between food tassium sorbate increases with increasing concentrations of
handling and food poisoning. An exception is the ovine bio- CO2 in packed atmospheres. Less is understood about the use
type, isolates of which also frequently produce enterotoxin. of preservatives to prevent toxin formation specifically. Nat-
PFGE has been used to type isolates from a meat processing urally occurring compounds, such as plant essential oils, are
plant and the results support the view that S. aureus on beef now also being considered as hurdles.
carcasses originate primarily from the hands of workers en- Predictive models, which model the growth rate of the
gaged in carcass dressing. organism given various input parameters such as pH and
The organism is generally regarded as a poor competitor temperature, have been reported to overpredict growth.
with other foodborne microorganisms. It, therefore, generally Therefore, they are generally, but not always, fail-safe. Other
does not grow and cause problems in raw meats. Because of its models predict the growth/no growth boundary in particular
tolerance to low water activity, its growth is favored in foods meat products. These may be useful tools for evaluating shelf
that are salted or dried, such as cured and fermented meats. It stability.
may also grow in foods such as cooked meats in which a In general, proper cleaning and sanitizing are effective in
competing flora has been reduced during processing. removing the organisms from manufacturing plants because
the organism is sensitive to sanitizers used in the food
industry.
Epidemiology

Staphylococcal intoxication contributes substantially to the

burden of illness acquired from contaminated food. In 2008, See also: Microbiological Analysis: DNA Methods; Indicator
in the EU, there were 291 outbreaks suspected as being Organisms in Meat; Standard Methods. Microbiological Safety of
attributable to staphylococcal intoxication, accounting for Meat: Bacillus cereus; Hurdle Technology. Microorganisms and
5.5% of the total number of outbreaks of foodborne disease/ Resistance to Antibiotics, the Ubiquity of: Antibiotic Resistance
intoxication. Of these outbreaks, 27.8% were verified. Two by Microorganisms. Modeling in Meat Science: Microbiology
patients died. It is generally thought that, as symptoms are
often self-limiting, only 10% of people with staphylococcal
intoxication seek medical treatment.
Typical outbreaks occur through postcooking contamin-
ation by food handlers, with subsequent temperature abuse of Further Reading
the food that allows growth of and toxin production by S.
aureus. For example, in an American outbreak where ham was Balaban, N., Rasooly, A., 2000. Staphylococcal enterotoxins. International Journal of
the implicated vehicle, contamination probably occurred Food Microbiology 61, 1–10.
when an infected food handler removed casings from the Cretenet, M., Even, S., Le Loir, Y., 2011. Unveiling Staphylococcus aureus
enterotoxin production in dairy products: A review of recent advances to face
hams which were subsequently stored at temperatures above new challenges. Dairy Science and Technology 91, 127–150.
10 °C for more than 15 h. De Buyser, M.L., Lombard, B., Schulten, S.M., et al., 2003. Validation of EN ISO
standard methods 6888 Part 1 and Part 2: 1999-Enumeration of coagulase-
positive staphylococci in foods. International Journal of Food Microbiology 83,
185–194.
Control and Preventative Measures International Commission on Microbiological Specifications for Foods, 1996.
Staphylococcus aureus. In: Roberts, T.A., Baird-Parker, A.C., Tompkin, R.B.
An important preventative measure centers around the (Eds.), Micro-organisms in Foods 5. Microbiological Specifications of Foodborne
training and personal hygiene of staff who handle food. Pathogens. London: Blackie Academic, pp. 299–333.
Staff should cover wounds with sticking plasters and gloves Johler, S., Stephan, R., 2010. Staphylococcal food poisoning: A current review.
Archiv für LebensmittelhygieneLebensmittelhyg 61, 197–236.
where appropriate, and refrain from touching their noses, Kluytmans, J.A.J.W., 2009. Methicillin-resistant Staphylococcus aureus in food
hair, or other parts of their body while working with food. products: Cause for concern or case for complacency? Clinical Microbiology and
If such events occur, then hands should be washed Infection 16, 11–15.
immediately. Lancette, G.A., Bennett, R.W., 2001. Staphylococcus aureus and staphylococcal
enterotoxins. In: Downes, F.P., Ito, K. (Eds.), Compendium of Methods for the
Because the organism will grow well on foods with a
Microbiological Examination of Foods, fourth ed. Washington, DC, USA:
background microflora reduced by cooking, it is essential to American Public Health Association, pp. 387–403.
avoid cross-contamination of cooked foods by bacteria from Seo, K.S., Bohach, G.A., 2007. Staphylococcus arueus. In: Doyle, M.P., Beuchat, L.
raw foods in which staphylococci are likely to be present. R., Montville, T.J. (Eds.), Food Microbiology: Fundamentals and Frontiers.
Holding foods at a temperature less than 7 °C will ensure that Washington, DC, USA: ASM Press, pp. 493–518.
Stewart, C.M., 2003. Staphylococcus aureus and staphylococcal enterotoxins. In:
the organism cannot grow and produce toxin. Hocking, A.D. (Ed.), Foodborne Microorganisms of Public Health Significance,
Product formulations can be manipulated to prevent the sixth ed. Sydney, Australia: AIFST (NSW Branch) Food Microbiology Group,
growth of the organism by using multiple ‘hurdles.’ For pp. 359–379.
 Microbiological Safety of Meat | Staphylococcus aureus 381

Relevant Websites http://www.fda.gov/Food/FoodSafety/FoodborneIllness/

FoodborneIllnessFoodbornePathogensNaturalToxins/BadBugBook/ucm070015.htm
http://www.aoac.org/testkits/tested methods.html FDA Bad bug book.
AOAC International. http://www.foodsafety.govt.nz/elibrary/industry/Staphylococcus_Aureus-
http://www.combase.cc/index.php/en/ Science_Research.pdf
ComBase. Foodborne pathogens data sheet: Staphylococcus aureus.
http://www.fda.gov/food/scienceresearch/laboratorymethods/ http://www.hc-sc.gc.ca/fn-an/alt_formats/hpfb-dgpsa/pdf/res-rech/mfhpb21-eng.pdf
bacteriologicalanalyticalmanualbam/ucm073674.htm Government of Canada method for the enumeration of S. aureus in foods.
FDA Bacteriological Analytical Manual: Staphylococcal Enterotoxins: Micro-slide http://pmp.arserrc.gov/PMPOnline.aspx
Double Diffusion and ELISA-based Methods. Predictive models: Pathogen modelling program (PMP) online.
http://www.fda.gov/Food/ScienceResearch/LaboratoryMethods/ http://www.textbookofbacteriology.net/staph.html
BacteriologicalAnalyticalManualBAM/ucm071429.htm Todar's online textbook of bacteriology.
FDA Bacteriological Analytical Manual: Staphylococcus aureus.