Alcoholic Fermentation / Second Edition, 1914

by Arthur Harden

Fermentation of Different Sugars by Yeast.

Many valuable ideas as to the nature of fermentation have been obtained by a consideration of the phenomena presented by the action of yeast on the different hexoses. Of these only glucose, fructose, mannose, and galactose are susceptible of alcoholic fermentation by yeast, the stereoisomeric hexoses prepared in the laboratory being unfermentable, as are also the pentoses, tetroses, and the alcohols corresponding to all the sugars. The yeast-cell is therefore much more limited in its power of producing fermentation than such an organism as, for example, Bacillus coli communis, which attacks substances as [p131] diverse as arabinose, glucose, glycerol and mannitol, and yields with all of them products of the same chemical character, although in varying proportions.

A careful examination of a number of different genera and species of the Saccharomycetaceæ and allied organisms by E. F. Armstrong [[1905]] has shown that all yeasts which ferment glucose also ferment fructose and mannose. Armstrong grew his yeasts in a nutrient solution containing the sugar to be investigated, and his experiments are open to the criticism that the organisms were hereby afforded an opportunity for becoming acclimatised to the sugar. His results, therefore, only demonstrate the fact that the organisms in question when cultivated in presence of the sugars examined brought about their fermentation, and do not exclude the possibility that the same organism when grown in presence of a different sugar might not be capable of fermenting the one to which it had in the other type of experiment become acclimatised.

This has actually been shown to be the case for galactose by Slator [[1908, 1]], and it is possible that this circumstance explains the negative results obtained by Lindner [[1905]] with S. exiguus and Schizosaccharomyces Pombe upon mannose, a sugar which, according to Armstrong, is fermented by both these organisms.

The same problem has been attacked quantitatively by Slator, who has shown that living yeast of various species and genera ferments glucose and fructose at approximately the same rate. Moreover, when the yeast is acted upon by various inhibiting agents, such as heat, iodine, alcohol, or alkalis, the crippled yeast also ferments glucose and fructose at the same rate.

With mannose the relations are somewhat different. The relative rate of fermentation of mannose and glucose by yeast is dependent on the variety of the yeast and the treatment which it has received. Fresh samples of yeast ferment mannose more quickly than glucose, but by older samples the glucose is the more rapidly decomposed. This is especially the case with yeast, the activity of which has been partly destroyed by heat, the relative fermenting power to mannose being sometimes reduced by this treatment from 120 per cent. of that of glucose to only 12 per cent. (Slator).

A further difference consists in the fact that with certain yeasts the rate of fermentation of glucose is somewhat increased by monosodium phosphate whilst that of mannose is unaffected [Euler and Lundeqvist, [1911]].

Mixtures of glucose and fructose are fermented by yeast at the [p132] same rate as either the glucose or the fructose contained in the mixture would be alone. When, however, mannose and glucose are fermented simultaneously interference between the reactions takes place, and this is especially evident when the yeast has comparatively little action on mannose. The following are the results obtained by Slator:—

Yeast.	Relative Rates.
	2·5 per cent. Glucose.	2·5 per cent. Mannose.	2·5 per cent. Glucose + 2·5 per cent. Mannose.
S. Thermantitonum	100	105	92
Brewery yeast, 53 per cent. activity destroyed by heat	100	21	33
Brewery yeast, 60 per cent. activity destroyed by heat	100	12	42

The case of galactose merits special attention. Previous investigations [see Lippmann, [1904], p. 734] have shown that the fermentation of galactose by yeast differs greatly from that of the other hexoses. The subject has been re-investigated by E. F. Armstrong [[1905]], and by Slator [[1908, 1]]. Armstrong carried out his experiments in the manner already described (p. [131]), and found that some yeasts had, and others had not, the power of fermenting galactose, although all were capable of fermenting glucose, fructose, and mannose.

Slator made quantitative experiments on the same subject. He was able to confirm the statement which had previously been made, that certain yeasts which have the property of fermenting galactose possess it only after the yeast has become acclimatised by culture in presence of the sugar. This was shown for brewery yeast and for the species mentioned below. This phenomenon is one of great interest and is strictly analogous to the adaptation of bacteria which has now been quite conclusively established [Neisser, [1906]].

Yeast.	Mode of Culture. Grown in:	Relative Rates.
		Glucose.	Galactose.
S. Carlsbergensis	wort	100	<1
"	hydrolysed lactose	100	86, 83, 85, 25, 46, 51, 69, 54, 155
S. Cerevisiæ	wort	100	<1
"	hydrolysed lactose	100	21, 26, 29
S. Thermantitonum.	wort	100	<1
"	hydrolysed lactose	100	77, 53, 35
S. Ludwigii	wort	100	<1
"	hydrolysed lactose	100	<1

[p133]

It will be seen that in one case the rate of fermentation of galactose was considerably greater than that of glucose. S. Ludwigii did not respond to the cultivation in hydrolysed lactose, but, as Slator points out, it is quite possible that repeated cultivation in this medium might effect the change, and this would be strictly analogous to the results obtained with bacteria. Slator's results have been confirmed by Harden and Norris, R. V. [[1910]], and by Euler and Johansson [[1912, 2]] who have made an exceedingly interesting study of the progress of the adaptation. As in the case of mannose the rates of fermentation of glucose and galactose are differently affected by agents such as heat and alcohol; moreover, the rate of fermentation of mixtures of dextrose and galactose is in no case either the sum or the mean of the rates obtained with the separate sugars. The temperature coefficient of the fermentation of galactose also differs slightly from that of the other hexoses.

Yeast.	Relative Rates.
	Glucose.	Galactose.	Glucose + Galactose.
S. Cerevisiæ	100	0	103
"	100	34	103
S. Carlsbergensis	100	155	119
S. Thermantitonum	100	76	124

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.

A third theory has also been suggested to explain these phenomena, according to which the various sugars are fermented by different enzymes [see Slator, [1908, 1]]. The uniformity of the result obtained with glucose and fructose suggests that these two sugars are fermented by the same enzyme (glucozymase), mannose and galactose by different ones (mannozymase and galactozymase). This would afford a simple explanation of the different rates of fermentation for different sugars and of different degrees of sensitiveness towards reagents.

If, however, a separate and independent mechanism were present for each sugar, the rate of fermentation of mixtures should be the sum of the rates for the constituents. This, as shown above, is not found to be the case, and it is therefore necessary to suppose, either that one sugar influences the fermentation of another in some unknown way, or that only a part of the mechanism of fermentation is specific for the particular sugar. Thus the enzyme may be specific and, the co-enzyme [p135] non-specific, so that only a certain maximum rate is attainable, or again, the supply of free phosphate may be the controlling factor.

In the prevailing state of ignorance as to the exact function of the co-enzyme and of the conditions upon which the velocity of fermentation in the cell depends, it is at present impossible to decide between these various theories, but they all offer points of attack which justify the hope that much further information can be obtained by experimental inquiry.

It will be seen from the foregoing that Buchner's discovery of zymase has opened a chapter in the history of alcoholic fermentation which is yet far from being completed. In every direction fresh problems present themselves, and it cannot be doubted that as in the past, the investigation of the action of the yeast-cell will still prove to be of fundamental importance for our knowledge of the mode in which chemical change is brought about by living organisms. [p136]

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INDEX.

• Acetaldehyde, as an intermediate product of alcoholic fermentation, [110].
  • — reduction of by yeast, [110].
• Acetone-yeast, [38].
• Alanine, as an intermediate product of alcoholic fermentation, [115].
• Alcohol, formation of, from sugar by alkalis, [97].
• Alcoholic fermentation, attempts to separate enzymes of, from yeast- cell, [15].
  • — — by-products of, [85].
  • — — equation of, [51].
  • — — Gay-Lussac's theory of, [4].
  • — — Iwanoff's theory of, [106].
  • — — kinetics of, [120], [128].
  • — — Lavoisier's views on, [3].
  • — — Liebig's theory of, [8].
  • — — Nägeli's theory of, [15].
  • — — of the amino-acids, [87].
  • — — — — theory of, [91].
  • — — Pasteur's researches on, [11].
  • — — Traube's enzyme theory of, [14].
• Alkalis, effect of, on hexoses, [96].
• Amino-acids, alcoholic fermentation of, [87].
  • — stereoisomerides of, fermented at different rates by yeast, [89].
• d-Amyl alcohol, formation of from isoleucine, [86].
• Antiprotease in yeast-juice, [42], [65].
• Antiseptics, action of, on yeast-juice, [19], [36].
• Arsenate, effect of, on fermentation by yeast-juice and zymin, [73].
  • — — on autofermentation of yeast-juice, [80].
  • — nature of acceleration produced by, [78].
• Arsenite, effect of, on fermentation by yeast-juice, [77].
  • — — on autofermentation of yeast-juice, [80].
  • — nature of acceleration produced by, [78].
• Autofermentation of yeast-juice, [33], [119].
  • — — effect of arsenates and arsenites on, [80].
• Baeyer's theory of fermentation, [99].
• Boiled yeast-juice, effect of, on fermentation by yeast-juice, [41].
• Carboxylase, [81], [93].
  • — relation of to alcoholic fermentation, [83].
• Co-enzyme, effect of electric current on, [67].
  • — enzymic destruction of, [63].
  • — of yeast-juice, [59].
  • — precipitation of, by ferric hydroxide, [67].
  • — properties of, [63].
  • — removal of, from yeast-juice, [59].
  • — separation from phosphate and hexosephosphate, [67].
• Concentration of sugar, effect of, on fermentation by yeast-juice, [34].
• Dauerhefe, [38].
• Diastatic enzyme of yeast-juice, [33].
• Dihydroxyacetone, fermentability of, [104].
  • — formation of, in fermentation, [105].
• Dried yeast (Lebedeff), [24], [38].
• Endotryptase, [20].
• Enzyme action, laws of, [121].
• Enzymes, combined with protoplasm, [126].
• Equation of alcoholic fermentation, [51].
• Fermentation by yeast-juice, causes of cessation of, [64].
• Fermenting complex, [63].
  • — power of yeast-juice, estimation of, [27].
• Formaldehyde, production of in alcoholic fermentation, [117].
• Formic acid theory of fermentation, [114].
• Fructose, fermentation of, by yeast-juice, [32].
  • — — in presence of phosphate, [73].
  • — relation of, to fermenting complex, [74].
• Fusel oil, formation of, from amino-acids, [85].
• Galactose, fermentation of, by yeast, [131].
  • — fermentation of, by yeast-juice, [32].
• Glucose, fermentation of, by yeast-juice, [32].
• Glyceraldehyde, fermentability of, [104].
• Glyceric acid, fermentation of, [108].
• Glycerol, formation in fermentation, [95].
• Glycogen as an intermediate product of alcoholic fermentation, [116].
  • — fermentation of, by yeast-juice, [33].
  • — removal of, from yeast, [39].
• Grinding of yeast by hand, [22].
  • — — — mechanical, [23].
• Glutamic acid, decomposition of, by yeast, [90].
• Hefanol, [38].
• Hexosediphosphoric acid phenylhydrazone, hydrazine salt of, [50].
• Hexosemonophosphoric acid osazone, hydrazine salt of, [50].
• Hexosephosphatase, [54].
• — effect of arsenate and arsenite on action of, [79].
• Hexosephosphate, constitution of, [51].
  • — enzymic decomposition of, in yeast-juice, [56].
  • — — hydrolysis of, [51].
  • — formation of, [48].
  • — hydrolysis of, by acids, [49].
  • — preparation of, [48].
  • — properties of, [49].
  • — theory of formation of, [57], [117],
• Hexoses, action of alkalis on, [96],
• Isoamyl alcohol, formation from leucine, [87].
• Isoleucine, decomposition of, by yeast, [87].
• α-Ketonic acids, fermentation of, [81].
• Lactic acid, destruction of, by yeast-juice, [102].
  • — — formation from sugars by alkalis, [97].
  • — — — of, in yeast-juice, [102].
  • — — non-fermentability of, by yeast, [103].
  • — — theory of fermentation, [102].
• Leucine, decomposition of, by yeast, [87].
• Maceration extract, preparation of, [25].
• Mannose, fermentation of, by yeast, [131].
  • — — of by yeast-juice, [32].
• Methylglyoxal, conversion of, into lactic acid, [101].
  • — non-fermentability of, [104].
  • — as an intermediate product of alcoholic fermentation, [113].
• Oxalacetic acid, formation of, from tartaric acid, [101].
• Permanent yeast, [38].
• Phenylethyl alcohol, [88].
• Phosphate, changes of, in alcoholic fermentation, [47].
  • — effect of, on fermentation by yeast-juice, [42].
  • — — — — — by zymin, [46].
  • — — — — — of fructose, [73].
  • — — of on total fermentation of yeast-juice, [54].
  • — influence on fermentation of concentration of, [71].
  • — inhibition by, [71].
• Phosphates, essential for alcoholic fermentation, [55].
• Proteoclastic enzyme of yeast, [20].
• Protoplasmic theory of activity of yeast-juice, [19].
• Pyruvic acid, fermentation of, [81].
  • — — theory of fermentation, [109].
• Rate of fermentation, controlling factors of, [119].
• Reductase, intervention of, in alcoholic fermentation, [111].
• Serum, effect of, on fermentation by yeast-juice, [41].
• Succinic acid, formation of, in fermentation, [89].
  • — — formed from glutamic acid by yeast, [90].
• Synthetic enzyme in yeast-juice, [32].
• Temperature coefficient of fermentation by yeast, [129].
  • — — — — by zymin, [122].
  • — — — esterification of phosphoric acid by yeast extract, [58].
• Tryptophol, [88].
• Tyrosol, [88].
• Wohl's theory of fermentation, [101].
• Yeast, action of toluene on, [124].
  • — and yeast-juice, fermentation by, compared, [29], [124].
  • — discovery of the vegetable nature of, [5].
  • — fermentation by, [127].
  • — — of different sugars by, [130].
  • — influence of concentration of dextrose on fermentation by, [128].
  • — — — — of, on rate of fermentation, [129].
  • — — of toluene on autofermentation of, [126].
  • — nature of the process of fermentation by, [123].
  • — temperature coefficient of fermentation by, [129].
  • — theories of fermentation by, [133].
• Yeast-juice and yeast, fermenting powers compared, [29], [124].
  • — co-enzyme of, [59].
  • — dialysis of, [59], [62].
  • — effect of arsenate on fermentation by, [75].
  • — — of concentration of sugar on fermentation by, [34].
  • — — of dilution on fermentation by, [35].
  • — — of phosphate on total fermentation, produced by, [54].
  • — estimation of fermenting power of, [27].
  • — evaporation of, [37].
  • — filtration of through gelatin filter, [59].
  • — precipitation of, [38].
  • — preparation of, [21].
  • — properties of, [19].
  • — ratio of alcohol and carbon dioxide, produced by, [30].
  • — synthesis of complex carbohydrate by, [31].
  • — variation of rate of fermentation by, with concentration of sugar, [121].
• Zymase, Buchner's discovery of, [16].
  • — enzymic destruction of, [64].
  • — properties of, [18].
  • — regeneration of inactive, [64].
  • — separation from co-enzyme, [59].
• Zymin, [21], [38].
  • — fermentation by, [39].
  • — rate of fermentation by, [39].
  • — temperature coefficient of fermentation by, [122].

ABERDEEN: THE UNIVERSITY PRESS

TRANSCRIBER'S NOTE:

With a few exceptions, original spelling and grammar were retained. The cover image is a modified version of the original scanned image from archive.org.

This book contains several uncommon Unicode characters, including "→", "↓", "⇌", "⇅", "│", "╱", "╲", "═", "║". A font and ebook reader software capable of rendering these is necessary for reading this book.

Full stops, middle dots "·", or even "˙"—Dot Above"—were variably (randomly?) used in the original as decimal points and for denoting chemical bonds. These have been rendered as middle dots herein.

The reference to Colin's paper on [page 5] has been changed from 1826 to 1825, to agree with the corresponding entry in the Bibliography. The reference to Turpin's paper on [page 8] was changed to 1838 from 1839, for the same reason.

Tables and formulas have been edited for clarity and readability, while honoring the original form. For example, the two sequential equations at the top of [page 110] originally had "+O" and "−H₂O" written under the two arrows, and the two equations appeared on one line. These have been converted into two equations on two lines, with the arrow subtext moved into the equations.

The incorrect formula for the enol II. in the equation for Glucose dehydration near bottom of [page 101] was corrected.