23.
Energy Transformations: Respiration and Photosynthesis
Glucose Metabolism: Overall Plan
As the diagram at the right indicates, all that is
obtained from the conversion of glucose to lactate is two
molecules of ATP, which makes anaerobic glycolysis a very inefficient
process. Yeasts in wine can get nineteen times as much energy per
mole of glucose by oxidizing it all the way to C02 and
H20, than by fermenting it anaerobically to ethanol.
The winemaker uses this fact to encourage rapid growth of the yeast
culture early in the wine-making process by bubbling air through
the crushed grapes. No ethanol is produced under these circumstances,
but the yeasts multiply rapidly in the presence of a large energy
supply. After the yeast colony is large, aeration is halted and
the grape juice in the vat is covered with a layer of carbon dioxide
to keep out oxygen. The yeasts stop multiplying, turn off their
citric acid cycle, and settle down to the anaerobic conversion of
glucose to ethanol - less rewarding for the yeast, but more rewarding
for the winemaker.
Bacteria have a much richer chemistry. All bacteria
begin with fermentation, and for some this is the end of the process.
They degrade glucose (and a few other molecules) anaerobically to
a number of different waste products; ethanol, or lactic, formic,
acetic, propionic, or butyric acids. Other bacteria respire using
02, giving off H20 as eucaryotes do. Still
others can use sulfate or nitrate as their oxidizing agents. Oxidation
with nitrate (yielding N2) appears to be a recent special
adaptation in some bacteria that always prefer 02 if
available. But sulfate respiration (yielding H2S) may
be an independent and very old line of metabolic evolution.