Models of cellular energetic and

metabolism
As we move beyond the compartmental models of the type
above, the level of metabolic details required increases.
At this point , it will be useful to consider the types of
reactions that occur within the cell. Metabolic pathway
can be distinguished as catabolic and anabolic. In
catabolism, energy-containing molecules, such as
carbohydrates, hydrocarbons and other reduced carboncontaining compounds are degraded to CO2 or other
oxidized end-products and the energy is stored in ATP,GTP,
and other energy-rich compounds. In anabolism,
intermediates and end-products formed from catabolism
are incorporated into cell (such as DNA, RNA, lipids,
carbohydrates, etc.) and their intermediate precursors
(amino acids, purines and pyrimidines, simple sugars
etc.). Anabolic reactions generally require energy, which is
supplied via ATP and other high-energy phosphates
generated during catabolism. As the concentration of
these high-energy intermediates within the cell is rather
small, anabolism is linked to catabolism and ATP is rapidly
turned over. This implies that energy producing and
energy consuming processes must be tightly regulated
within the cell. It is thus necessary to consider both
carbon and energy flows within the cell in developing
these more complex models. An example of such a model
is given in the following section.

A model for aerobic growth of the

yeast saccharomyces cevisiae

The yeast s.cevisiae (bakers yeast) has been well studied

and considerable information on its cell cycle. Regulation
and metabolism is known. Hall and co-workers have
formulated a model of the rather complex metabolism
exhibited by s.cerevisae when grown on glucose. This
yeast can use either the respiratory pathway , resulting in
the formation of ethanol, carbon dioxide and cell mass .at
low growth rates. metabolism is fully oxidative ,i.e., the
respiratory quotient(RQ) defined as the ratio of the rates
of carbon dioxide production to oxygen consumption ,is
unity and Y x/s is 0.50gm cells/gm glucose.this situation
is maintained up to a critical growth rate .beyond which
the metabolism becomes increasingly fermentative .in the
fermentative pathway,the yield coefficient decreases and
there n an increase in the specific carbon dioxide
production rate (Qco2)and ethanol production . this
critical growth rate is, slightly higher than the value of
max on ethanol. there is change in the enzyme pattern
that reflects this switch from respiration to fermentation:
typical respiratory enzymes ,such as isocitrate lyase,
malate dehydrogenase and the cytochromes are
repressed at high growth rates ,and glycolysis provides
the main source of energy .at low growth rates.reduced
level of glycolytic enzymes are found.
As the growth rate increases , the percentage of budding
yeast cells increases almost linearly. using this linear
relationship and the mean generation time (ln2/), the
length of the budding period can be calculated. There is
little variation in the duration of the budding period at
different growth rates. At low growth rates, the
generation time increase due to lengthening of the gapphase following cell division (the post-mitotic gap, or G1
phase). Thus, referring to the cell cycle, the time periods
for DNA replication (S), mitosis (G2) and cell division (M)

phases are all constant. The duration of the G1 phase

appears to be variable. During the single-cell G1 phase,
substrate is accumulated and there is a buildup of reserve
carbohydrates (trehalose and glycogen) within the cell
that are then depleted for energy and carbon during the
period of budding.
The model we shali examine is based on this two-stage
breakdown of the cell cycle. The length of the G1 phase
depends on the availability of the limiting substrate; the
length of the division phase (the sum of G2, M and S
phases) is assumed to be independent of substrate. The
cell mass is considered to be comprised of two parts: A
mass, which carried out substrate uptake and energy
production; and B mass, which carries out reproduction
and division. B mass is converted to A mass at a constant
rate, whereas A mass consumes substrate and produces B
mass at a variable rate.
We now turn to the regulation of respiration and
fermentation. The repression of respiratory enzymes by
glucose (the crabtree effect) was initially thought to
result from glucose acting as a catabolite repressor. more
recent evidence suggests that a high catabolic flux is the
direct cause of respiratory inhibition and that glucose
concentration plays a secondary role. Thus the model
proposes that both glycolysis and respiration are carried
out by A mass and both provide energy for growth.