Alcoholic Fermentation: Second Edition, 1914Harden, Arthur
Science
Alcoholic Fermentation: Second Edition, 1914
Harden, Arthur
Fermentation
(1) 2C{6}H{12}O{6} + 2PO{4}HR{3} =
2CO{2} + 2C{2}H{6}O + 2H{2}O + C{6}H{10}O{4}(PO{4}R{2}){2}.
According to this, two molecules of sugar are concerned in the change,
the carbon dioxide and alcohol being equal in weight to one [p052]
half of the sugar used, and the hexosephosphate and water representing
the other half.
Additional confirmation of this equation is afforded by the
determination of the ratio between sugar used and carbon dioxide
evolved when a known weight of sugar together with an excess of
phosphate is added to yeast-juice at 25°. The phenomena then observed
are precisely similar to those which occur when a phosphate is
added to a fermenting mixture of yeast-juice and excess of sugar as
described above. The rate of fermentation rapidly rises and then
gradually falls until a rate is attained approximately equal to that
of the autofermentation of the juice in presence of phosphate. At this
point it is found that the extra amount of carbon dioxide evolved,
beyond that which would have been given off in the absence of added
sugar, bears the ratio expressed in equation (1) to the sugar added
[Harden and Young, 1910, 2]. The results of four estimations made in
this way were (/a/) 0·2 grams of glucose gave 26·5 and 27·9 c.c. of
carbon dioxide at N.T.P.; (/b/) 0·2 grams of fructose gave 27·9 and
28·9 c.c. The carbon dioxide calculated from the sugar added in each
of the four cases is 26·96 c.c.
It has also been shown by Euler and Johansson [1913] that in the
fermentation of a mixture of equivalent amounts of phosphate and
glucose, the whole of the glucose had disappeared when the whole of
the phosphate had become esterified.
CYCLE OF CHANGES UNDERGONE BY PHOSPHATE IN ALCOHOLIC FERMENTATION.
According to equation (1) the free phosphate present is used up in the
reaction, and the question at once arises whether there is any source
from which a constant supply of free phosphate can be elaborated
in the juice, or whether some other change occurs which results in
the formation of carbon dioxide and alcohol in the absence of free
phosphate. The experimental evidence points in the direction of the
former of these alternatives, but the question is a very difficult one
to decide with absolute certainty.
When a mixture of a phosphate with yeast-juice and sugar is examined
at intervals and the amount of free phosphate estimated, the following
stages are observed:--
1. During the initial period of accelerated fermentation following the
addition of the phosphate, the concentration of free phosphate rapidly
diminishes.
2. At the close of this period, the amount of free phosphate [p053]
present is very low, and, as long as active fermentation continues, no
marked increase in it occurs.
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