Alcoholic Fermentation: Second Edition, 1914Harden, Arthur
Science
Alcoholic Fermentation: Second Edition, 1914
Harden, Arthur
Fermentation
Assuming that his conclusion that all yeasts which ferment glucose
also ferment fructose and mannose is correct, Armstrong has drawn
attention to the fact that these three hexoses are also related by the
possession of a common enolic form (p. 97) and has suggested that this
enolic form is the substance actually fermented to carbon dioxide and
alcohol [1904].
The idea that such an intermediate form is the direct subject of
fermentation has much to recommend it. In the first place it is
almost certain, as already pointed out, that the sugars in aqueous
solution do exist, although to a very small extent, in this enolic
form. The slow rate at which equilibrium is established in aqueous
solution, however, must be taken as definite evidence that under
these circumstances the enolic form is only produced very slowly
[compare Lowry, 1903]. This has been used by Slator [1908, 1] as an
argument against the probability of the preliminary conversion of
the sugars into the enolic form before fermentation. It appears,
however, quite possible that under the influence of the fermenting
complex of the yeast-cell, or of special enzymes, this change might
occur much more rapidly, [p134] and at different rates with the
different sugars. This reaction might in fact control the observed
rate of fermentation. This conception affords a simple explanation of
the different rates of fermentation of mannose and glucose, and also
of galactose, the enolic form of which is quite different, by yeast
under different circumstances, but does not explain the uniformity
of rate observed by Slator for glucose and fructose nor the results
with mixtures of sugars. The direct fermentation of a common enolic
form is also consistent with the fact that the same hexosephosphate is
produced from all three hexoses.
Slator himself prefers the view that the first stage of fermentation
consists in the rapid combination of the sugar with the enzyme,
producing a compound, which then breaks up at a rate which determines
the observed rate of fermentation. This rate will of course vary
with the nature of the compound, so that if two sugars form the
same compound they will be fermented at the same rate; if they form
different compounds, different rates may result. Slator supposes
that glucose and fructose form the same compound with the enzyme.
This, however, appears to involve an intramolecular change of the
same order as the production of the enolic form, and moreover is not
absolutely essential, as it is probably sufficient to suppose that
the two compounds derived from glucose and fructose are very similar,
although possibly not absolutely identical. Mannose and galactose, on
the other hand, form stereoisomeric compounds, and the capacity of the
fermenting complex to form these compounds may be affected by various
agents to a different extent from its capacity for combining with
glucose or fructose.
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