Alcoholic Fermentation: Second Edition, 1914Harden, Arthur
Science
Alcoholic Fermentation: Second Edition, 1914
Harden, Arthur
Fermentation
Precisely the same products are given with phenylhydrazine by the
hexosephosphoric acid prepared from glucose, mannose, and fructose,
proving that all these sugars yield the same hexosediphosphoric acid,
a point of fundamental importance.
Direct measurements of the molecular weight of the acid by the
freezing-point method, combined with the determination of the degree
of dissociation by the rate of cane-sugar inversion, are indecisive,
but indicate that the acid has a molecular weight considerably higher
than that required for a triosephosphoric acid.
A similar uncertainty attaches to the determination of the molecular
weight from the freezing-point depression and conductivity of the
acid potassium salt [Euler and Fodor, 1911]. Euler however concludes
[p051] that both a hexosediphosphoric acid and a triosemonophosphoric
acid are formed, but has not prepared any derivatives of the latter.
As regards the constitution of the hexosephosphoric ester several
suggestions have been made by Young, but no decisive evidence at
present exists. The identity of the products from glucose, mannose,
and fructose may be explained by regarding the acid as a derivative
of the enolic form common to these three sugars (p. 97), or by
supposing that portions of two sugar molecules may be concerned in its
production. The formation and composition of the hydrazone and osazone
are of great importance as they indicate that in all probability
one of the phosphoric acid residues is united with the carbon atom
adjacent to the carbonyl group of the hexose. They moreover render it
certain that the original phosphoric ester is a hexosediphosphoric
ester and not a triosemonophosphoric ester.
Hexosediphosphoric acid has not as yet been discovered in the animal
body. The action of a number of enzymes upon it has been examined
[Euler, 1912, 2; Euler and Funke, 1912; Harding, 1912; Plimmer, 1913]
with the following results.
The lipase of castor oil seeds, a glycerol extract of the intestinal
mucous membrane of the rabbit and pig, and an aqueous extract of bran
have a slow hydrolytic action, whereas pepsin and trypsin are without
effect. Feeding experiments with rabbits and dogs indicate that the
ester is capable of hydrolysis in the animal body, a large proportion
of the phosphorus being excreted as inorganic phosphate. The ester is
also decomposed by /Bacillus coli communis/.
It is remarkable that the hexosephosphate is not fermented nor
hydrolysed by living yeast, a fact observed by Iwanoff, Harden and
Young, and Euler, although, according to the experiments of Paine
[1911], the yeast cell is at all events partially permeable to the
sodium salt.
THE EQUATION OF ALCOHOLIC FERMENTATION.
An equation can readily be constructed for the reaction in which
hexosephosphate is formed, the data available being the formula of the
product and the relation between the phosphate added and the carbon
dioxide and alcohol produced:--
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