Food Microbiology (CFD20203)

Chapter 2 : Factors That Influence Microbial Growth

CHAPTER 2: FACTORS THAT INFLUENCE MICROBIAL GROWTH

Objectives:

After this lesson student should be able to:

1. Describe factors that influence microbiological growth
a. Intrinsic factors
- pH, moisture content, redox potential, nutrient content,
antimicrobial constituents of the food, and biological structures.
b. Extrinsic factors
- temperature, relative humidity, concentration of gases, and the
presence of other microbes.
2. State the interaction between growth factors
3. Used the factors to control the microbiological growth in food products

INTRODUCTION

Since microbes are living organisms, they must have certain things to continue with life.
The main item that they must have is a source of energy. However, like all living
organisms, there must also be specific environmental parameters, which must be met in
order for microbes to survive. And like other differing organisms, these parameters are
different for different organisms to survive on foods. There are intrinsic parameters of the
food itself. These are factors that are exhibited by the food or surface on which the
microbe is growing. There are also extrinsic factors that affect the growth of microbes.
These are factors that are not part of the food or surface that the microbe is growing on.

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General Principles of Microbial Growth & survival

Microbial growth generally proceeds through a series of phases:

• Lag phase: time for microorganisms to become accustomed to their new

environment. There is little or no growth during this phase.
• Log phase: microorganisms logarithmic, or exponential, growth begins; the rate
of multiplication is the most rapid and constant.
• Stationary phase: the rate of multiplication slows down due to lack of nutrients
and build-up of toxins. At the same time, microorganisms are constantly dying so
the numbers actually remain constant.
• Death phase: cell numbers decrease as growth stops and existing cells die off

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Factors that affect the microbial growth can be categorized as extrinsic and intrinsic
factors
Intrinsic factors are those that are characteristic of the food itself
Extrinsic factors are those refer to the environment surrounding the food.

2.1 Intrinsic factors

There are six intrinsic factors that affect the growth and survival of microbes on food.
They are:

i) pH
ii) moisture content
iii) nutrient content
iv) antimicrobial constituents of the food
v) biological structures
vi) redox potential

2.1.1 pH

Increasing the acidity of foods, either through fermentation or the addition of weak acids,
has been used as a preservation method since ancient times. In their natural state, most
foods such as meat, fish, and vegetables are slightly acidic while most fruits are
moderately acidic. A few foods such as egg white are alkaline.

The pH of a substance is its relative acidity or alkalinity. It is read on a scale of 1 to 14

with pH 7 being neutral. As one moves toward pH 1 the substance is more acidic, while if
one move toward pH 14, the substance becomes more alkaline.

The idea of acidity or alkalinity is based on ability of a substance to accept or donate

protons. If a substance donates protons (H+ ions), then it is considered acidic. If the
substance accepts protons, then it is alkaline. Another way to look at pH is whether a

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substance donates or accepts electrons. By this way of thinking, if a substance accepts

electrons, then it is acidic. However, if it donates electrons, then it is considered alkaline.
Most microbes grow well when the pH is around 7, or neutral.

Microorganisms can be placed in one of the following groups based on their optimum pH
requirements:

Acidophiles
pH optimum between 2 and 6
Neutrophiles
pH Optimum between 6 and 8
Alkalophiles
pH optimum between 8 and 11

Different microbes can grow at different pH ranges. This is shown in the following table.

Organism pH Range
Molds 0.2 - 11
Yeasts 1.5 - 8.5
Salmonella 3.6 - 9.5
Listeria monocytogenes 4.2 - 9.6
Yesinia entercolitica 4.2 - 9.0
Escherichia coli 4.3 - 9.0
Clostridium botulinum 4.3 - 8.5
Bacillus cereus 5.0 - 9.5
Campylobacter 5.0 - 9.0
Shigella 5.0 - 9.2
Vibrio parahaemolyticus 5.0 - 11
Vibrio cholerae 5.0 - 9.5
Clostridium perfringens 5.0 - 8.5

Table adapted from Modern Food Microbiology, 6th Ed., by James M. Jay, p.36

• Most spoilage bacteria grow best near neutral pH

• Pathogenic bacteria even more narrow in tolerance range of near neutral

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• Yeast and molds have much greater tolerance to acidic (lower) pH

• The optimum pH range is usually quite narrow so that small changes in the pH
can have large effects on the growth rate of the organism
• On either end of pH range organisms will experience increased lag and
generation times
• NaCl can act as antagonist as “optimal concentrations” extending growth range
of an organism (at pH) but if added in excess of optimum level will begin to act
synergistically

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2.1.2 Moisture

Microorganism needs water in available form to grow in food product. The control of the
moisture content in foods is one of the oldest exploited preservation strategies. Food
microbiologist generally describes the water requirements of microorganisms in term of
water activity (aw).
Water activity is defined as the ratio of water vapor pressure of the substrate to the vapor
pressure of pure water at the same temperature.

aw = p/po

p= Vapor pressure of the solution

po =Vapor pressure of the solvent
aw for pure water is 1.00

However, vapor pressures are hard to calculate, so another method has been devised for
measuring the water activity of a substance. To do this uses the following equation:

Aw = moles of water ÷ (moles of water + moles of solute)

By this method, pure water has a water activity of 1.00. However if 1 mole of a solute is
added, then the solution has a water activity of 0.98. Water activity is measured on a
scale of 0.00 to 1.00. Most fresh foods have a water activity of 0.99. Most spoilage
microbes cannot survive if the water activity is below 0.91. However, there are some
yeasts and molds that can survive at water activity of 0.61.

Most flesh meat, vegetable and fruits have aw values that are close to growth level of
most microorganism ( 0.97 – 0.99)

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Bacteria have optimum growth at aw near to 1

Microorganism aw minimum
Most bacteria 0.91
Most yeast 0.88
Most Mold 0.80
Halophile bacteria 0.75
Xerophile mold 0.65
Osmophile yeast 0.60

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2.1.3 Nutritional requirements

Microorganisms require certain basic nutrients for growth and maintenance of metabolic
functions. The amount and type of nutrients required range widely depending on the
microorganism. These nutrients include water, a source of energy, nitrogen, vitamins, and
minerals.

Foodborne microorganisms can derive energy from carbohydrates, alcohols, and amino
acids. Most microorganisms will metabolize simple sugars such as glucose. Others can
metabolize more complex carbohydrates, such as starch or cellulose found in plant foods,
or glycogen found in muscle foods. Some microorganisms can use fats as an energy
source.

In addition to a proper physical environment, microorganisms also depend on an

available source of chemical nutrients. Microorganisms are often grouped according to
their energy source and their source of carbon.

A. Energy source

1. Phototrophs use radiant energy (light) as their primary energy source.

2. Chemotrophs use the oxidation and reduction of chemical compounds as their

primary energy source.

B. Carbon source

Carbon is the structural backbone of the organic compounds that make up a living cell.
Based on their source of carbon bacteria can be classified as autotrophs or heterotrophs.

1. Autotrophs: require only carbon dioxide as a carbon source. An autotroph can

synthesize organic molecules from inorganic nutrients.

2. Heterotrophs: require organic forms of carbon. A Heterotroph cannot synthesize

organic molecules from inorganic nutrients.

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Combining their nutritional patterns, all organisms in nature can be placed into one of
four separate groups:

1. photoautotrophs
2. photoheterotrophs
3. chemoautotrophs
4. chemoheterotrophs.

1. Photoautotrophs use light as an energy source and carbon dioxide as their main
carbon source. They include photosynthetic bacteria (green sulfur bacteria, purple sulfur
bacteria, and cyanobacteria), algae, and green plants. Photoautotrophs transform carbon
dioxide and water into carbohydrates and oxygen gas through photosynthesis.

Cyanobacteria, as well as algae and green plants, use hydrogen atoms from water to
reduce carbon dioxide to form carbohydrates, and during this process oxygen gas is given
off (an oxygenic process). Other photosynthetic bacteria (the green sulfur bacteria and
purple sulfur bacteria) carry out an anoxygenic process, using sulfur, sulfur compounds
or hydrogen gas to reduce carbon dioxide and form organic compounds.

2. Photoheterotrophs use light as an energy source but cannot convert carbon dioxide
into energy. Instead they use organic compounds as a carbon source. They include the
green nonsulfur bacteria and the purple nonsulfur bacteria.

3. Chemoautotrophs use inorganic compounds such as hydrogen sulfide, sulfur,

ammonia, nitrites, hydrogen gas, or iron as an energy source and carbon dioxide as their
main carbon source.

4. Chemooganoheterotrophs use organic compounds as both an energy source and a

carbon source. Saprophytes live on dead organic matter while parasites get their
nutrients from a living host. Most bacteria, and all protozoans, fungi, and animals are
chemoorganoheterotrophs.

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C. Nitrogen source

Nitrogen is needed for the synthesis of such molecules as amino acids, DNA, RNA and
ATP. ATP is Adenosine triphosphate which is a compound containing the nitrogenous
base adenine, the carbohydrate ribose, and three phosphates. Breakdown of ATP provides
the energy needed to carry out energy-requiring processes in cells. Depending on the
organism, nitrogen, nitrates, ammonia, or organic nitrogen compounds may be used as a
nitrogen source.

D. Minerals

1. sulfur

Sulfur is needed to synthesisize sulfur-containing amino acids and certain vitamins.

Depending on the organism, sulfates, hydrogen sulfide, or sulfur-containing amino acids
may be used as a sulfur sorce.

2. phosphorus

Phosphorus is needed to synthesize phospholipids, DNA, RNA, and ATP. Phospholipids

are the building block of cellular membranes. Phospholipids are composed of phosphate,
glycerol, and fatty acids. The phosphate-glycerol portion of the molecule is water soluble
(polar) while the fatty acid portion is water insoluble (non-polar). Phosphate ions are the
primary source of phosphorus.

3. potassium, magnesium, and calcium

These are required for certain enzymes to function as well as additional functions.

4. iron

Iron is a part of certain enzymes.

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5. trace elements

Trace elements are elements required in very minute amounts, and like potassium,
magnesium, calcium, and iron, they usually function as cofactors (Organic molecules or
ions that assist many enzymes in their reactions) in enzyme reactions. They include
sodium, zinc, copper, molybdenum, manganese, and cobalt ions. Cofactors usually
function as electron donors or electron acceptors during enzyme reactions.

e. Water

f. Growth factors

Growth factors are organic compounds such as amino acids, purines ( Double-ringed
nitrogenous bases found in DNA and RNA. They include adenine and guanine.),
pyrimidines ( Single-ringed nitrogenous bases found in DNA and RNA. They include
thymine, cytosine, and uracil), and vitamins that a cell must have for growth but cannot
synthesize itself. Organisms having complex nutritional requirements and needing many
growth factors are said to be fastidious ( Having complex nutritional requirements and
needing many growth factors.).

The microorganisms that usually predominate in foods are those that can most easily
utilize the nutrients present. Generally, the simple carbohydrates and amino acids are
utilized first, followed by the more complex forms of these nutrients. The complexity of
foods in general is such that several microorganisms can be growing in a food at the same
time. The rate of growth is limited by the availability of essential nutrients.

The abundance of nutrients in most foods is sufficient to support the growth of a wide
range of foodborne pathogens. Thus, it is very difficult and impractical to predict the
pathogen growth or toxin production based on the nutrient composition of the food.

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2.1.5 Antimicrobial constituent of food

Antimicrobial constituents are naturally occurring substances within the food that provide
some activity against the growth of microbes. These constituents can be as varied as the
microbes and food substrates themselves. Some examples include:

1. spices and essential oils that possess antimicrobial activities

o eugenol (cloves, cinnamon, sage), cinnamic aldehyde (cinnamon),

thymol (sage, oregano), isothymol (sage), allicin (garlic), allyl
isothiocyanate (mustard),

2. enzymes

o lysozyme in eggs and milk

3. lactoperoxidase system in milk

o enzyme lactoperoxidase, H2O2, thiocyanate (pseudomonas very

sensitive)
o thiocyanate and H2O2 levels must be enhanced to attain activity-
can be used where refrigeration is uncommon- will extend shelf
life of raw milk for a few days

2.1.6 Biological structures

Plant and animal derived foods, especially in the raw state, have biological structures that
may prevent the entry and growth of pathogenic microorganisms. Examples of such
physical barriers include skin of fruits and vegetables, shell of nuts, animal hide, egg
cuticle, shell, and membranes. Milk, however, is a fluid product with no barriers to the
spreading of microorganisms throughout the product.

Plant and animal foods may have pathogenic microorganisms attached to the surface or
trapped within surface folds or crevices. Intact biological structures thus can be important

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in preventing entry and subsequent growth of microorganisms. Several factors may

influence penetration of these barriers. The maturity of plant foods will influence the
effectiveness of the protective barriers. Physical damage due to handling during harvest,
transport, or storage, as well as invasion of insects can allow the penetration of
microorganisms.

During the preparation of foods, processes such as slicing, chopping, grinding, and
shucking will destroy the physical barriers. Thus, the interior of the food can become
contaminated and growth can occur depending on the intrinsic properties of the food. For
example, Salmonella spp. have been shown to grow on the interior of portions of cut
cantaloupe, watermelon, honeydew melons given sufficient time and temperature.

The egg is another good example of an effective biological structure that, when intact,
will prevent external microbial contamination of the perishable yolk. For the interior of
an egg to become contaminated by microorganisms on the surface, there must be
penetration of the shell and its membranes. In addition, the egg white contains
antimicrobial factors. When there are cracks through the inner membrane of the egg,
microorganisms penetrate into the egg. Factors such as temperature of storage, relative
humidity, age of eggs, and level of surface contamination will influence internalization.
For example, conditions such as high humidity and wet and dirty shells, along with a
drop in the storage temperature will increase the likelihood for entry of bacteria. If eggs
are washed, the wash water should be 12 oC (22 oF) higher than the temperature of the
eggs to prevent microbial penetration. After washing, the eggs should be dried and then
cooled. The Food and Drug Administration (FDA) published a final rule that applies to
shell eggs that have not be processed to destroy all live Salmonella before distribution to
the consumer.

Heating of food as well as other types of processing will break down protective biological
structures and alter such factors as pH and aw. These changes could potentially allow the
growth of microbial pathogens.

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2.1.6 Redox potential

The redox potential of a substance is the ease by which it gains or loses electrons. If a
substance loses electrons, it is described as being oxidized. If a substance gains electrons,
it is said to be reduced. When electrons move they create a potential difference, which
can be measured.

In regard to microbes, each has a specific tolerance or need for an amount of oxygen in
its environment. Oxidized environments provide aerobic conditions. These oxidized
environments have a redox potential that is measured in positive millivolts. Reduced
environments provide for an anaerobic environment. Their redox potential is measured in
negative millivolts.

As you might suspect, those microbes that are aerobic in nature need an environment that
has a positive redox potential, whereas anaerobes need an environment with a negative
redox potential. Those microbes that are facultative in nature can survive in environments
with varying redox potentials. Since pH can also be measure by using the loss or
acceptance of electrons, the redox potential of a substance is dependent on the pH of that
substance. So, when the redox potential of a substance is give, it should be given with a
pH. Most redox potentials are measured at pH 7.0

Redox potential (Eh) requirements and minimum tolerances differ for different species
but in general

aerobes will not tolerate low numbers (require positive value)

facultatives and anaerobes can boh tolerate numbers in the same low
range (negative values)
but the anaerobes will not start to grow until the redox has reached a
critical maximum which is much lower than the facultatives
plant foods tend to have positive values (300-400mV)
minced meats (200mV)
-aerobic bacteria and mold spoil these

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solid meats have low numbers (-200mV)—after rigor mortis (before are
in +range and inhibit anaerobes)
cheeses vary in the negative range –20—200mV (microaerophillic,
facultatives found here)
also by products of metabolism can further decrease the Eh (H2S will
lower Eh )

Available Oxygen: Microorganisms can be classified according to their oxygen

requirements necessary for growth and survival:

• Obligate Aerobes: oxygen required

• Facultative: grow in the presence or absence of oxygen
• Microaerophilic: grow best at very low levels of oxygen
• Aerotolerant Anaerobes: oxygen not required for growth but not harmful if
present
• Obligate Anaerobes: grow only in complete absence of oxygen; if present it can
be lethal

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2.2 Extrinsic factors

Along with the six intrinsic factors that affect microbial growth, there are extrinsic factors
that will also have an effect on the ability of microbes to grow and survive in their
environment. These include:

a. temperature
b. relative humidity
c. concentration of gases
d. presence of other microbes.
e. Chemical

2.2.1 Temperature
Each type of microorganism has upper and lower temperature limit. Beyond which they
are not be able to grow. Between these upper and lower limits, there is an optimum
temperature at which growth is maximized.
There is a wide variation in the temperature limits and optima for different
microorganism. There are generally three categories: psychrophiles, mesophiles, and
thermophiles.

Psychrophiles

Psychrophiles are organisms that have an optimum temperature range of

20 - 30°C, but can survive at temperature below 7°C. Some of these organisms
actually prefer the lower end of the range. An example of a psychrophiles
includes Psedomonas spp. Other examples are :

• Pseudomonas, Acinetobacter, Psychrobacter, and Enterococcus members

are important bacterial spoilage psychrophiles
• yeasts- Rhodotortula, Candida, Saccharomyces
• molds- Mucor, Penicillium, Rhizopus,

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• Clostridium botulinum E, Listeria monocytogenes, Aeromonas hydrophila,

Yersinia enterocolitica, B. cereus are exapmles of pathogenic
psychrophiles

Mesophiles

Mesophiles are organisms that have an optimum growth temperature between 30 -

40°C, but can survive at temperatures between 20 - 45°C and minimum at 5-10oC.
An example of a mesophile is Staphylococcus aureus.

Thermophiles

Thermophiles are organisms that grow well above 25-45oC and have an optimum
temperature range between 55 - 65°C, but can survive at temperatures
approaching 85°C. Examples of thermophiles include Bacillus and Clostridium
genera.

Thermotolerants is an extreme thermophiles- Thermus genus

Some endospore producing strains are thermotolerant after spore production

some sporeformers true thermophiles capable of withstanding high temps for
periods of time and will grow at between 40-60oC

o Bacillus sternothermophilus
o Bacillus coagulans
o Clostridium thermosaccharolyticum

are some examples of microbes which can be a problem in the canning industry-
survive the processing and then in warmer climates will proceed to spoil the
canned food.

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2.2.2 Relative Humidity

The relative humidity of the environment surrounding the food is important because it
affects the water activity of the food. If the relative humidity is low, moisture may be
drawn from the food into the surrounding air to produce equilibrium, thus reducing the
water activity.

if food has low water activity it will need a storage condition of low humidity in
order to maintain that low aw at the surface of the product

Foods will attempt to reach equilibrium with environment- so will pick up water
from the environment if of a lower aw

foods with high aw will lose moisture to low humidity environment

Generally, foods that depend on certain aw for safety or shelf life considerations will need
to be stored such that the environment does not markedly change this characteristic.
Thus, processors and distributors need to provide for appropriate storage conditions to
account for this fact.
Packaging, as discussed previously in this chapter, will play a major role in the
vulnerability of the food to the influence of relative humidity. But even within a sealed
container, moisture migration and the phenomenon of environmental temperature

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fluctuation may play a role. It has been observed that certain foods with low aw can be
subject to moisture condensing on the surface due to wide environmental temperature
shifts. This surface water will result in microenvironments favorable to growth of
spoilage, and possibly pathogenic, microorganisms.

As a general guideline, the product should be held such that environmental moisture,
including that within the package, does not have an opportunity to alter the aw of the
product in an unfavorable way.

2.2.3 Concentration of gases

The concentration of gases in the environment has a specific effect on the types of
microbes that can grow in the environment. Aerobic organisms require oxygen in
amounts above 12%. Some organisms, like Campylobacter, require minimal oxygen
(5%). Oxygen can be lethal to anaerobes. Some organisms require carbon dioxide in
certain levels.

Gases inhibit microorganisms by two mechanisms. First, they can have a direct toxic
effect that can inhibit growth and proliferation. Carbon dioxide (CO2), ozone (O3), and
oxygen (O2) are gases that are directly toxic to certain microorganisms. This inhibitory
mechanism is dependent upon the chemical and physical properties of the gas and its
interaction with the aqueous and lipid phases of the food. Oxidizing radicals generated by
O3 and O2 are highly toxic to anaerobic bacteria and can have an inhibitory effect on
aerobes depending on their concentration. Carbon dioxide is effective against obligate
aerobes and at high levels can deter other microorganisms. A second inhibitory
mechanism is achieved by modifying the gas composition, which has indirect inhibitory
effects by altering the ecology of the microbial environment. When the atmosphere is
altered, the competitive environment is also altered. Atmospheres that have a negative
effect on the growth of one particular microorganism may promote the growth of another.
This effect may have positive or negative consequences depending upon the native
pathogenic microflora and their substrate. Nitrogen replacement of oxygen is an example
of this indirect antimicrobial activity

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Altering gas composition of environment can help retard surface spoilage without
reducing the humidity at which a product is stored
Increasing CO2 (to 10%), could controlled or modified the atmosphere

can retard the spoilage of foods by molds -fruits/vegetables

increase shelf life of meats (esp. vacuum-packed)

Much work has been done on investigating the effects of different gas mixtures on a
product. The optimal effect will be achieved by trial and error- finding a similar situation
in the literature, trying it and adjusting it to your product.

2.2.4 Presence of other microbes

All microbes need food to survive. If conditions are optimum for one microbe, but less
than optimum for another microbe on the same food, the first microbe will have more
success. And all microbes produce waste products. The waste from one microbe may be
detrimental to another microbe living in the same area. This is what as known as
competitive inhibition. One microbe can excrete substances that will inhibit the growth of
another microbe. Of course, there is also symbiosis. In symbiosis, two or more organisms
share the same food supply. One or all of the organisms usually excretes a substance that
can be used by another microbe for its benefit.

Some organisms can produce inhibitory substances are better competitors for nutrient
supply. For example: -
- bacteriocins (lactic acid bacteria) - chemical compounds produced by microorganisms
that inhibit other microorganisms (example is nisin)
-antibiotics-secondary metabolites produced by microorganisms that inhibit or kill a
wide spectrum of microorganisms

For both mechanisms mentioned above- background population (the inhibitors) must be
larger (viable cells) than the competitor strain.

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2.2.5 Chemical : Salt, additive

Chemicals whether from outside or purposely added into the food also can influence the
microbial growth, thus increasing its ability in producing their own metabolites. The
effect of the chemicals towards toxin producer microorganisms become the interest of
study by the scientists because there are lots of chemicals especially preservative agents
have been used to extent the shelf life of the food.

The effect of 6 different food preservatives; propionic acids, benzoic acids, citric acids,
sodium acetate, sodium metabisulphite, and sodium chloride on the ability of
Aspergilulus parasiticus in aflatoxin production has been studied. It was found that
propionic acid at 0.1 % concentration in the SMKY is able to retard mycelium growth,
and aflatoxin production. The same result also was show by citric acids when the
concentration achieve 0.5 or 1.0%.

Due to the higher number of types and amounts of pesticides available and being used in
agriculture sector, the study of the effect of pesticides ( insecticides, fungicides,
herbicides and etc) towards microorganisms growth are very few. However, for example,
Phosphine and carbon disulphite (CS2), a type of pesticides widely used in cereals storage
warehouse, was found been able to increase the aflatoxin content inside the wheats by
Aspergillus flavus.

REFERENCES :
Ray B., (1996), Fundamental Food Microbiology, CRC Press, USA

Montville TJ, Matthews KR.(2001). Chapter 2: Principles which influence microbial

growth, survival, and death in foods. In: Doyle MP, Beuchat LR, Montville TJ, editors.
Food microbiology: fundamentals and frontiers. Washington (DC)

Mossel, D. et al. (1996). Essential of the Microbiology of Foods. John Wiley & Sons.

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