STOICHIOMETRY OF

MICROBIAL GROWTH AND Berces, Mark Gieronne B.

Berces, Ruemhel D.
Bunao, Jay-ar
PRODUCT FORMATION Vargas, Mark Gerard
 𝑴
Overall Growth Yield Coefficient 𝒀𝑿/𝑺
• the maximum yield of cell mass per unit mass of
substrate consumed when no maintenance is
considered.
• there are two other yield and maintenance coefficients
of importance are related to ATP consumption and
oxygen.
ATP Yield Coefficient 𝒀𝑿/𝑨𝑻𝑷
• represents the amount of biomass synthesized per mole
of ATP generated.

1 1 𝑚𝐴𝑇𝑃
𝐴𝑃 = 𝑀 +
𝑌𝑋/𝐴𝑇𝑃 𝑌𝑋/𝐴𝑇𝑃 𝐷

𝐴𝑃
Where 𝑌𝑋/𝐴𝑇𝑃 is the “apparent” yield of biomass
and 𝑚𝐴𝑇𝑃 is the rate of ATP consumption for maintenance
energy.
• Similarly, yields based on oxygen consumption can be
defined and calculated.

1 1 𝑚𝑂2
𝐴𝑃 = 𝑀 +
𝑌𝑋/𝑂2
𝑌𝑋/𝑂2 𝐷

Values of 𝑌𝑋/𝑂2 can vary from 0.17 to 1.5 g

biomass/g O2, depending on substrate and organism.
Respiratory Quotient (RQ)
• the moles of 𝐶𝑂2 produced per mole of oxygen
consumed.
• value provides an indication of metabolic state (for
example, aerobic growth versus ethanol fermentation
in baker’s yeast) and can be used in process control.
P/O ratio
• the ratio of phosphate bonds formed per unit of
oxygen consumed (g mole P/g atom O)
• indicates the efficiency of conversion of reducing
power into high-energy phosphate bonds in the
respiratory chain.
Proton/Oxygen Ratio (H/O)
• is the number of H+ ions released per unit of oxygen
consumed.
• Electron generation is directly related to proton
release.
• The generation of electrons results in the expulsion of
H+ that can be used directly to drive the transport of
some substrates or to generate ATP.
Regularities
• The complexity of mass and energy balances for cellular
growth can be decreased greatly through the recognition
that some parameters are nearly the same irrespective of
the species or substrate involved. These parameters can
be referred to as regularities.
• Three important regularities
• 26.95 kcal/g equivalent of available electrons transferred to
oxygen (coefficient of variation of 4%),
• 4.291 g equivalent of available electrons per quantity of biomass
containing 1 g atom carbon, and
• 0.462 g carbon in biomass per gram of dry biomass. It has also
been observed that 𝑌𝑋/𝑒 − = 3.14 ± 0.11 g dry wt/g equivalent of
electrons.
TOICHIOMETRIC CALCULATIONS

ELEMENTAL BALANCES
ELEMENTAL BALANCES
• It is a material balance for biological reactions, the composition of
the substrates, products, and cellular materials must be known.
• Aim to balance all elements of reactant side to product side (law
of conservation of matter)
• Provide no insight into the energetics of a bioreaction.
• No extracellular products other than water and carbon dioxide.
• CH1.8O0.5N0.2, a typical cellular composition; 1 mole of biological
material is define as the amount containing 1 gram atom of
carbon, CHαOβNδ .

CHmOn + a O2 + b NH3 → c CHα Oβ Nδ + d H2O + e CO2

Simple Elemental Balance
CHmOn + a O2 + b NH3 → c CHα Oβ Nδ + d H2O + e CO2

C:1=c+e H: m + 3b = cα + 2d
O: n + 2a = cβ + d + 2e
N: b = cδ
Respiratory Quotient
Moles of CO2
𝑒 formed
RQ =
𝑎 Moles of O2
formed

• The ratio of the carbon dioxide produced over

the oxygen consumed by an organism, tissue, or
cell in a given time.
• Varies for different metabolic fuels or substrates
which reflects net substrates oxidation.
 Degree of Reduction
The number of equivalents of available electrons per
gram atom C
The degree of reduction of any element in a compound
is Equal to the valence of this element.
The degrees of reduction for some key elements are :
C=4, H=1, N=-3, O=-2, P=5, S=6
Example on how to calculate
degree of reduction

A high degree of reduction indicates a low

degree of oxidation
Consider the aerobic production of a single
extracellular product:

The degree of reduction of substrate, biomass and

product are:

Note that CO2, H2O and NH3 the degree of reduction is zero
For such a data set, we would typically choose a carbon, a
nitrogen and an available-electron balance.
• Carbon Balance • Available-Electron Balance
𝑐+𝑑+𝑓 =1 (Eq 7.10) 𝑐𝛾𝑏 + 𝑑𝛾𝑝 = 𝛾𝑠 − 4𝑎 (Eq 7.12)

• Nitrogen Balance:
𝑐𝞭 + 𝑑𝑧 = 𝑏 (Eq 7.11)

With partial experimental data, it is possible to solve

this set of equation for example: RQ, Yx/p and Yx/s.
 𝑓
𝑅𝑄 =
𝑎
𝑐 = 𝑌𝑋Τ𝑆 (on a molar basis)
𝑑 = 𝑌𝑃Τ𝑆 (on a molar basis)

• An energy balance for an aerobic growth

𝑄0 𝑐𝛾𝑏 + 𝑄0 𝑑𝛾𝑝 = 𝑄0 𝛾𝑠 − 𝑄0 4𝑎 (Eq. 7.13)
𝑄0 = heat evolved per equivalent of available electrons
transferred to Oxygen
𝑄0 = 95.26 kcal/g of available electrons transferred to
Oxygen
• Equation 7.12 can be written as

𝒄𝜸𝒃 𝒅𝜸𝒑 𝟒𝒂
1= + + (Eq. 7.14a)
𝜸𝒔 𝜸𝒔 𝜸𝒔

1 = 𝝃𝒃 + 𝝃𝒑 + 𝜺 (Eq. 7.14b)

𝜺 = fraction of available electrons in the organic substrate that is

transferred to oxygen
𝝃𝒃 = fraction of available electrons that is incorporated to biomass
𝝃𝒑 = fraction of available electrons that is incorporated to
extracellular products
THEORETICAL PREDICTIONS OF
YIELD COEFFICIENTS
THEORETICAL PREDICTIONS OF YIELD COEFFICIENTS
Cell mass and product formation by
microorganisms can be described quantitatively by
yield coefficients expressed as the mass of cells or
product formed per unit mass of substrate
consumed, Yx/s, and Yp/s, for cells and product,
respectively.
Let's consider the overall
stoichiometric equation
for growth and
production:

where S, carbon source; N,

nitrogen source; X , cell mass;
P, product and s, n, o, p, w, e The cell mass yield coefficient and
are stoichiometric
coefficients. the product yield coefficient are

respectively, where Mx, Mp and Ms, are the molecular

weights of cell mass, product, and carbon source.
In aerobic fermentations,
the growth yield per
available electron in oxygen
molecules is approximately
3.14 ± 0.11 gdw(gram dry
weight) cells/electron when
ammonia is used as the
nitrogen source.
• The ATP yield (YX/ATP) in
many anaerobic
fermentations is
approximately 10.5 ± 2 gdw
cells/mol ATP.
• In aerobic fermentations,
this yield varies between 6
and 29. When the energy
yield of a metabolic pathway
is known (N moles of ATP
produced per gram of
substrate consumed), the
growth yield YX/S can be
calculated using the
following equation:
Example:
Estimate the theoretical growth and product yield coefficients for
ethanol fermentation by S. cerevisiae as described by the following
overall reaction:

Solution:
Since YX/ATP= approx. 10.5 gdw/mol ATP and since glycolysis yields
2 ATP/mol of glucose in yeast,
• For complete conversion of glucose to ethanol by the yeast
pathway, the maximal yield would be

• while for CO2 the maximum yield is

Cell Composition
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