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Communications to the Editor

Yield Coefficients for Cell Mass and

Product Formation
J. Hong
Biochemical Engineering Program, School of Engineering, University of
California, lrvine, California 92717
Accepted for publication, March 1, 1988

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/,and Yp/,for cells and product, respectively. With the yield coefficients, the material balance equations for cells, substrate, and product can be
straightforwardly formulated. However, it is frequently observed in the literature that the yield coefficients are misused, resulting in incorrect contradictory material balance
equations to the definition of yield coefficients. The objective of this communication is to clarify this point.
Let's consider the overall stoichiometric equation for
growth and production:

sS + nN

+ 00,

+ pP + wH,O + eCO,



The substrate consumption is determined as:

As can be seen from eqs. (4), ( 5 ) , and (6) the substrate

consumption or product formation rate is not independent
of the cell accumulation rate. This is due to the assumption
that the overall metabolic activity of microorganisms can
be described by the single stoichiometry, eq. (1).
The following extended version of the mass balance equation are also used for certain cases of product formation:

where S, carbon source; N , nitrogen source; X , cell mass;

P, product and s, n, 0,p . w ,e are stoichiometric coefficients. The theoretical yield coefficients can be determined
from the above stoichiometry with known chemical formula for S, N , X and P.
The cell mass yield coefficient and the product yield coefficient are
= -Mx





= -PMP



respectively, where M,, M p and M , are the molecular

weights of cell mass, product, and carbon source. The
usual manner of calculating the yield coefficients is to
measure the mass of cells or product produced and substrate consumed. If the fermentation is carried out in a
constant volume batch fermentor, the mass balance equations are:

= px


Biotechnology and Bioengineering, Vol. 33, Pp. 506-507 (1989)

0 1989 John Wiley & Sons, Inc.


-dt= w





where m is the maintenance coefficient expressed as a substrate demand to maintain cell viability per unit of cells per
time, and qp is the specific rate of product formation.
These equations are formulated based upon the assumption
that the substrate consumption rate is determined from the
following three parallel reactions:

+ n , N + 0,0,--+

+ w,H,O + elC02


+ w,H,O + e,C02


for the cell growth, and


+ n 2 N + 0,0,

for the product formation,

S + n3N +


Maintenance + w,H,O

+ e3C02

for the maintenance.

CCC 0006-3592/89/040506-02$04.00

The specific conversion rate of eq. (10) is p and that of

eq. (1 1) is qp. The specific conversion rate for the maintenance is m. Since the formulation of eq. (9) is based upon
the parallel conversion stoichiometry equations, eqs. (lo),
(Il), and (12), the definitions of Yi/sand Y&, in eq. (9) are
absolutely different from those in eqs. (5) and (6). The cell
mass yield coefficient, Y:/s in eq. (9) is the cell mass produced per mass of substrate consumed only for the conversion (10).

substrate consumed over some period of time, these yield

coefficients should not be used in eq. (9). If so, the substrate consumption will be erroneously counted twice. To
evaluate Yi/sin eq. (9), it is frequently assumed in the literature that no substrate loss is associated with the product
formation, then Yd,s can be stoichiometrically set from
eq. (11). For example, in the case of ethanol production
from glucose, the theoretical yield of Yils is determined to
be 0.51 g of ethanol/g of glucose based upon the following stoichiometry:

The product yield coefficient, Y i / sin eq. (9), is the product

formed per mass of substrate consumed only for the conversion (1 I),

Therefore, if the cell mass and the product are formed

simultaneously for a growth associated product formation,
experimental determination of Y:,s and Yi/s is not straightforward. The fraction of carbon source consumed for
eq. (10) and that for eq. (11) should be known. Also the
fraction of carbon source consumed for cell maintenance
should be known. However, the experimental determination of these fractions is not an easy task, especially when
the cell mass and the product are formed simultaneously. If
the yield coefficients are determined in the usual manner
by measuring the mass of cells or product produced and

-+ 2C,H,OH

+ 2C0,


Since ethanol is the major product during the alcoholic fermentation by yeast, the above stoichiometry is a reasonable approximation to a real one. However, if the product
is a minor component, in other words, the fermentation is
associated with production of other components, a simple
formulation of stoichiometry such as eq. (15) involves a
drastic assumption which should be validated experimentally. In summary, the yield coefficients determined in the
usual manner by measuring the mass of cells or product
produced and substrate consumed should not be used in the
mass balance equations of the form, eqs. (7), (8), and (9).
The validity of the extended mass balance equation,
eq. (9), should be examined by checking whether the stoichiometry involved in the product formation is a physically
reasonable representation of metabolic activity. If the stoichiometry in the product formation can not be determined
accurately, the extended mass balance equation, eq. (9),
should not be used.