The effect is not produced when the concentration of the phosphate is so high that the rate of fermentation of fructose is itself greatly lowered.
This remarkable inductive effect is specific to fructose and is not produced when glucose is added to mannose or fructose, or by mannose when added to glucose or fructose, under the proper conditions of concentration of phosphate in each case.
This interesting property of fructose, taken in connection with the [p075] facts that this sugar in presence of phosphate is much more rapidly fermented than glucose or mannose, and that the optimum concentration of phosphate for fructose is much higher than for glucose or mannose, appears to indicate that fructose when added to yeast-juice does not merely act as a substance to be fermented, but in addition, bears some specific relation to the fermenting complex.
All the phenomena observed are, indeed, consistent with the supposition that fructose actually forms a permanent part of the fermenting complex, and that, when the concentration of this sugar in the yeast-juice is increased, a greater quantity of the complex is formed. As the result of this increase in the concentration of the active catalytic agent, the yeast-juice would be capable of bringing about the reaction with sugar in presence of phosphate at a higher rate, and at the same time the optimum concentration of phosphate would become greater, exactly as is observed. The question whether, as suggested above, fructose actually forms part of the fermenting complex, and the further questions, whether, if so, it is an essential constituent, or whether it can be replaced by glucose or mannose with formation of a less active complex, remain at present undecided, and cannot profitably be more fully discussed until further information is available.
It must, moreover, be remembered that different samples of yeast-juice vary to a considerable extent in their relative behaviour to glucose and fructose, so that the phenomena under discussion may be expected to vary with the nature and past history of the yeast employed.
IV. Effect of Arsenates on the Fermentation of Sugars by Yeast-Juice and Zymin.
The close analogy which exists between the chemical functions of phosphorus and arsenic lends some interest to the examination of the action of sodium arsenate upon a mixture of yeast-juice and sugar, and experiments reveal the fact that arsenates produce a very considerable acceleration in the rate of fermentation of such a mixture [Harden and Young, [1906, 3]; [1911, 1]]. The phenomena observed, however, differ markedly from those which accompany the action of phosphate.
The acceleration produced is of the same order of magnitude as that obtained with phosphate, but it is maintained without alteration for a considerable period, so that there is no equivalence between the amount of arsenate added and the extra amount of fermentation effected. Further, no organic arsenic compound corresponding in composition with the hexosephosphates appears to be formed.
Increase of concentration of arsenate produces a rapid inhibition of [p076] fermentation, probably due to some secondary effect on the fermenting complex, possibly to be interpreted as the formation of compounds incapable of combining with sugar and hence unable to carry on the process of fermentation. An optimum concentration of arsenate therefore exists just as of phosphate, at which the maximum rate is observed, and this optimum concentration and the corresponding rate vary with different samples of juice and are less for glucose than for fructose. The rate of fermentation by zymin is relatively less increased than that by yeast-juice.
Owing to the fact that the rate is permanently maintained the addition of a suitable amount of arsenate increases the total fermentation produced to a much greater extent than phosphate.