With the formation of succinate, the two big energy-releasing and
C02-producing steps of the cycle are over, and the original
six-carbon citrate has been degraded to a four-carbon molecule.
However, more energy is still available. Succinate is oxidized to
fumarate with the storage of energy in FADH2, fumarate
is rearranged to malate, and malate finally is oxidized to oxaloacetate
with the simultaneous reduction of NAD+. The cycle is
completed when oxaloacetate combines with acetyl CoA and another
turn of the wheel begins.
The unfinished business is the machinery for reoxidizing NADH and
FADH2 and making use of their energy. This is the topic
of the next section. At this point we can stop and draw a balance
sheet of the entire energy situation, from glycolysis through the
citric acid cycle.

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This is equivalent to a total of (4 X 1) + (10 X 3) + (2 X 2) =
38 ATP molecules per molecule of glucose. Of the total 686 kcal
released per mole of glucose, 38 X 7.3 = 277 kcal are saved, a 40%
overall efficiency. The other 409 kcal are not entirely useless.
They ensure the thermodynamic spontaneity of the reaction, and provide
body heat:

Several energy-producing pathways besides glycolysis funnel together
and enter the citric acid cycle to produce energy. When fats are
used as an energy source, the fatty acids are broken down into two-carbon
acetate and fed into the cycle. During the metabolism of proteins,
some amino acids are converted into pyruvate or acetate and then
enter the cycle. Thus the biochemical machinery that probably evolved
to make maximum use of the products of glycolysis now is used with
many other processes. Any molecule that can be broken down to acetate
can enter the citric acid cycle and yield energy.
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