pH is a measure of the hydrogen ion activity of a solution. pH is defined as the negative logarithm of the hydrogen ion concentration (expressed in terms of molarity).
pH
= -log [H+] = log(1/[H+])
The pH scale extends from
pH 0.0 to pH 14.0.
The habitats in which
microorganisms grow vary widely-from pH 0 to 2 at the acidic end to alkaline
lakes and soil that may have pH values between 9 and 10.
pH strongly affects
microbial growth. Each species has a definite pH growth range and pH growth
optimum.
- Acidophiles have their growth optimum between pH 0 and 5.5;
- Neutrophiles, between pH 5.5 and 8.0;
- Alkalophiles prefer the pH range of 8.0 to 11.5.
- Extreme alkalophiles have growth optima at pH 10 or higher.
In general, different
microbial groups have characteristic pH preferences. Most bacteria and
protists (protozoa and algae) are neutrophiles. Most fungi prefer more acidic
surroundings, about pH 4 to 6; photosynthetic protists also seem to favour
slight acidity.
Many archaea are
acidophiles. For example, the archaeon Sulfolobus acidocaldarius is a
common inhabitant of acidic hot springs; it grows well around pH 1 to 3 and at
high temperatures. The archaea Ferroplasma acidarmanus and Picrophilus
oshimae can grow at pH 0, or very close to it.
Although microorganisms
will often grow over wide ranges of pH and far from their optima, there are
limits to their tolerance
Maintenance of pH
Drastic variations in
cytoplasmic pH can harm microorganisms by disrupting the plasma membrane or
inhibiting the activity of enzymes and membrane transport proteins. Most
procaryotes die if the internal pH drops much below 5.0 to 5.5. Changes
in the external pH also might alter the ionization of nutrient molecules and
thus reduce their availability to the organism.
Microorganisms respond to
external pH changes using mechanisms that maintain a neutral
cytoplasmic pH. Several mechanisms for adjusting to small changes in external
pH have been proposed.
Ø The
plasma membrane is impermeable to protons.
Ø Neutrophiles
have an antiport transport system to exchange potassium for protons.
Ø Extreme
alkalophiles like Bacillus alcalophilus maintain their internal pH
closer to neutrality by exchanging internal sodium ions for external protons.
Ø Internal
buffering also may contribute to pH homeostasis.
- If
the external pH becomes too acidic, other mechanisms come into play.
ü When
the pH drops below about 5.5 to 6.0, Salmonella enterica serovar
Typhimurium and E. coli synthesize an array of new proteins as
part of their acidic tolerance response (ATR).
ü A
proton-translocating ATPase is activated and makes more ATP or pump protons out of the cell and thus contributes to this protective
response,
ü If
the external pH decreases to 4.5 or lower, chaperone proteins such as acid
shock proteins and heat shock proteins are synthesized. These
prevent the acid denaturation of proteins and aid in the refolding of denatured
proteins.
Microorganisms frequently change the pH of their own habitat by producing acidic or basic metabolic waste products. Fermentative microorganisms form organic acids from carbohydrates, whereas chemolithotrophs like Thiobacillus oxidize reduced sulfur components to sulfuric acid. Other microorganisms make their environment more alkaline by generating ammonia through amino acid degradation. Because microorganisms change the pH of their surroundings, buffers often are included in media to prevent growth inhibition by large pH changes.
Phosphate is a commonly used buffer and is a good example of buffering by a weak acid (dihydrogen phosphate, H2PO4-) and its conjugate base (monohydrogen phosphate, HPO4 2-).
H+ + HPO4 2-⎯⎯→
H2PO4-
OH- + H2PO4- ⎯⎯→ HPO4 2-
+ HOH
If protons are added to
the mixture, they combine with the salt form to yield a weak acid. An increase
in alkalinity is resisted because the weak acid will neutralize hydroxyl ions
through proton donation to give water.
Peptides and amino acids in complex media also have a strong buffering effect.
pH |
[H+] Molarity |
|
Environmental examples |
Microbial
examples |
|
0 |
10–0
|
Increasing
acidity
|
Concentrated
nitric acid |
Ferroplasma,
Picrophilus oshimae |
|
1 |
10–1
|
Gastric
contents, acid thermal springs |
Dunaliella
acidophila |
|
|
2 |
10–2
|
Lemon
juice Acid mine drainage |
Cyanidium
caldarium, Thiobacillus thiooxidans, Sulfolobus
acidocaldarius |
|
|
3 |
10–3
|
Vinegar,
ginger ale Pineapple |
|
|
|
4 |
10–4
|
Tomatoes,
orange juice Very acid soil |
|
|
|
5 |
10–5 |
Cheese,
cabbage Bread |
Physarum
polycephalum, Acanthamoeba castellanii |
|
|
6 |
10–6 |
Beef,
chicken Rain water Milk, Saliva |
Lactobacillus acidophilus, E.coli, Pseudomonas aeruginosa, Euglena gracilis, Paramecium bursaria |
|
|
7 |
10–7 |
Neutrality |
Pure
water, Blood |
Staphyloccus
aureus |
|
8 |
10–8 |
Increasing
alkalinity |
Seawater |
Nitrosomonas
spp. |
|
9 |
10–9 |
Strongly
alkaline soil, Alkaline lakes |
|
|
|
10
|
10–10 |
Soap |
Microcystis
aeruginosa, Bacillus alcalophilus |
|
|
11
|
10–11 |
Household
ammonia |
|
|
|
12
|
10–12 |
Saturated
calcium hydroxide solution |
|
|
|
13
|
10–13 |
Bleach,
Drain opener
|
|
|
|
14 |
10–14
|
|
|
|
The pH scale and microorganisms with their growth optima.
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