Alcoholic Fermentation: Second Edition, 1914Harden, Arthur
Science
Alcoholic Fermentation: Second Edition, 1914
Harden, Arthur
Fermentation
When 0·1 gram of dihydroxyacetone in 5 c.c. of water was brought in
contact with 1 gram of living yeast, about half was fermented, 17 c.c.
of carbon dioxide (at 20° and 600 mm.) being evolved in excess of the
autofermentation of the yeast (13 c.c.). A much greater effect was
obtained by the aid of yeast-juice, and the remarkable observation
was made that whilst yeast-juice alone produced comparatively little
action a mixture of yeast-juice and boiled yeast-juice was much
more effective, quantities of 20 to 50 c.c. of yeast-juice mixed
with an equal volume of boiled juice, which in some experiments was
concentrated, yielding with 0·4, 1, and 2 grams of dihydroxyacetone
almost the theoretical amount of carbon dioxide and alcohol in
excess of that evolved in the absence of this substance. It was
further observed that the fermentation of this substance commenced
much more slowly than that of glucose. No explanation of either of
these facts has at present been offered. The conclusion drawn from
their experiments by Buchner and Meisenheimer that dihydroxyacetone
is readily fermentable, was confirmed by Lebedeff [1911, 1], who
further made the important observation that during the fermentation
of dihydroxyacetone the same hexosephosphoric acid is produced as is
formed during the fermentation of the hexoses. Lebedeff accordingly
propounded a scheme of alcoholic fermentation according to which the
hexose [p106] was first converted into two molecules of triose,
the latter being first esterified to triosephosphoric acid and
then condensed to hexosediphosphoric acid, which then underwent
fermentation, after being hydrolysed to phosphoric acid, and some
unidentified substance, probably an unstable modification of a hexose,
much more readily attacked by an appropriate enzyme than the original
glucose or fructose [1911, 1, pp. 2941-2].
The idea that the sugar is first converted into triose and this into
triosemonophosphoric acid had been previously suggested by Iwanoff
who postulated the agency of a special enzyme termed /synthease/
[1909, 1], and supposed that this triosemonophosphoric acid was then
directly fermented to alcohol, carbon dioxide and phosphoric acid.
According both to Iwanoff and Lebedeff the phosphoric ester is an
intermediate product and its decomposition provides this sole source
of carbon dioxide and alcohol. This is quite inconsistent with the
facts recounted above (Chap. III), which prove that the formation
of the hexosephosphate is /accompanied/ by an amount of alcoholic
fermentation exactly equivalent to the quantity of hexosephosphate
produced, and that the rate of fermentation rapidly falls as soon as
the free phosphate has disappeared, in spite of the fact that at that
moment the concentration of the hexosephosphate is at its highest,
whereas according to Iwanoff's theory it is precisely under these
conditions that the maximum rate of fermentation should be maintained.
Public-domain text, read in full here on John Shaqi.
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