Alcoholic Fermentation: Second Edition, 1914Harden, Arthur
Science
Alcoholic Fermentation: Second Edition, 1914
Harden, Arthur
Fermentation
Experiments with low concentrations of sugar are difficult to
interpret, the influence of the hydrolysis of glycogen and of
dextrins on the one hand, and the synthesis of sugar to more complex
carbohydrates on the other (p. 31), having a relatively great effect
on the concentration of the sugar. Unpublished experiments (Harden
and Young) indicate, however, that the velocity of fermentation
remains approximately constant, until a certain very low limit of
sugar concentration is reached, and then falls rapidly. The fall in
rate, however, only continues over a small interval of concentration,
after which the velocity again becomes approximately constant and
equal to the rate of autofermentation. During this last phase, as
already indicated, the velocity is generally controlled by the rate
of production of sugar and no longer by that of phosphate, this
substance being now present in excess. In other words, the rate of
fermentation of sugar by yeast-juice and zymin is not proportional
to the concentration of the sugar present as required by the law
of mass, but, after a certain low limit of sugar concentration, is
independent of this and is actually slightly decreased by increase in
the concentration of the sugar.
The relations here are very similar to those shown to exist by Duclaux
[1899] and Adrian Brown [1902] for the action of invertase on cane
sugar and are probably to be explained in the manner suggested by
the latter. According to this investigator, the enzyme unites with
the fermentable material, or as it is now termed, the substrate or
zymolyte, forming a compound which only slowly decomposes so that it
remains in existence for a perceptible interval of time. The rate of
fermentation depends on the rate of decomposition of this compound
and hence varies with its concentration. This conception leads to
the result that the rate of fermentation will increase with the
concentration of the substrate up to a certain limit and will then
remain [p122] constant, unless interfered with by secondary actions.
This limit of concentration is that at which there is just sufficient
of the material in question present to combine with practically
the whole of the enzyme, so that no further increase in its amount
can cause a corresponding increase in the quantity of its compound
with the enzyme or in the rate of fermentation which depends on the
concentration of that compound.
The curve relating the rate of action of such an enzyme with the
concentration of the zymolyte therefore consists of two portions, one
in which the rate at any moment is proportional to the concentration
of the zymolyte, according to the well-known law of the action
of mass, and a second in which the rate at any moment is almost
independent of that concentration, approximately equal amounts being
decomposed in equal times whatever the concentration of the substrate.
Public-domain text, read in full here on John Shaqi.
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