CHAPTER IX.

THE MECHANISM OF FERMENTATION.

The analysis of the process of alcoholic fermentation by yeast-juice

and other preparations from yeast which has been carried out in the

preceding chapters has shown that the phenomenon is one of a very

complex character. The principal substances directly concerned in the

change appear to be the enzyme and co-enzyme of the juice, a second

enzyme, hexosephosphatase, and, in addition, sugar, phosphate, and

the hexosephosphate formed from these. During autofermentation two

other factors are involved, the complex carbohydrates of the juice,

including glycogen and dextrins, and the diastatic ferment by which

these are converted into fermentable sugars. It is also possible that

the supply of free phosphate is partially provided by the action of

proteoclastic ferments on phosphoproteins. Under special circumstances

the rate at which fermentation proceeds may be controlled by the

available amount of any one of these numerous substances.

When the juice from well-washed yeast is incubated, the phenomenon

of autofermentation is observed. The juice contains an abundant

supply of enzyme, co-enzyme, and phosphate or hexosephosphate, and

in this case the controlling factor is usually the supply of sugar,

which is conditioned by the concentration of the diastatic enzyme or

of the complex carbohydrates as the case may be. When this is the

case the measured rate of fermentation is the rate at which sugar is

being produced in the juice, this being the slowest of the various

reactions which are proceeding under these circumstances. If sugar be

now added, an entirely different state of affairs is set up. As soon

as any accumulated phosphate has been converted into hexosephosphate,

the normal rate of fermentation which is usually higher than that

of autofermentation is attained, and, provided that excess of sugar

be present, fermentation continues for a considerable period at a

slowly diminishing rate and finally ceases. During the first part of

this fermentation the rate is controlled entirely by the supply of

free phosphate, and this depends mainly on the concentration of the

hexosephosphatase and of the hexosephosphate, and only in a secondary

degree on the decomposition [p120] of other phosphorus compounds by

other enzymes and on the concentration of the sugar. The amount of

hexosephosphate in yeast-juice is usually such that an increase in

its concentration does not greatly affect the rate of fermentation,

and hence the measured rate during this period represents the rate

at which hexosephosphate is being decomposed, and this in its turn

depends on the concentration of hexosephosphatase, which is therefore

the controlling factor. As fermentation proceeds, the concentration

of both enzyme and co-enzyme steadily diminishes, as already

explained, probably owing to the action of other enzymes, so that at

an advanced stage of the fermentation, the controlling factor may

be the concentration of either of these, or the product of the two

concentrations (see p. 122). The hexosephosphatase appears invariably

to outlast the enzyme and co-enzyme. The condition at any moment

could be determined experimentally if it were possible to add enzyme,

co-enzyme and hexosephosphatase at will and so ascertain which of

these produced an acceleration of the rate.

Unfortunately this can at present be only very imperfectly

accomplished, owing to the impossibility of separating these

substances from each other and from accompanying matter which

interferes with the interpretation of the result.

A third condition can also be established by adding to the fermenting

mixture of the juice and sugar a solution of phosphate. The supply

of phosphate is now almost independent of the action of the

hexosephosphatase, and the measured rate represents the rate at which

reaction (1), p. 51, can occur between sugar and phosphate in the

presence of the fermenting complex consisting of enzyme and co-enzyme.

This change is controlled, so long as sugar and phosphate are present

in the proper amounts, by the concentration of the fermenting complex

or possibly of either the enzyme or the co-enzyme. If only a single

addition of a small quantity of phosphate be made, the rate falls as

soon as the whole of this has been converted into hexosephosphate and

the reaction then passes into the stage just considered, in which the

rate is controlled by the production of free phosphate.

Although these varying reactions have not yet been exhaustively

studied from the kinetic point of view, owing to the experimental

difficulties to which allusion has already been made, investigations

have nevertheless been carried out on the effect of the variation of

concentration of yeast-juice and zymin as a whole, as well as of the

carbohydrate. Herzog [1902, 1904] has made experiments of this kind

with zymin, and Euler [1905] with yeast-juice, whilst many of the

results [p121] obtained by Buchner and by Harden and Young are also

available.

The actual observations made by these authors show that the initial

velocity of fermentation is almost independent of the concentration

of sugar within certain limits, but decreases slowly as the

concentration increases. When the velocity constant is calculated on

the assumption that the reaction is monomolecular [see Bayliss, 1914,

Chap. VI], approximate constancy is found for the first period of the

fermentation. This method of dealing with the results is, however,

as pointed out by Slator, misleading, the apparent agreement with

the law of monomolecular reactions being probably due to the gradual

destruction of the fermenting complex.

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.

The results of the experiments with yeast-juice therefore indicate

that what is being measured is a typical enzyme action, but afford

no information as to which of the many possible actions is the

controlling one, a fact which must be ascertained for each particular

case in the manner indicated above.

Clowes [1909], using washed zymin free from fermenting power

and adding various volumes of boiled yeast extract, found that

the velocity of reaction was proportional to the product of the

concentrations of zymin and yeast extract up to a certain optimum

concentration. He interprets these concentrations as representing

the concentrations of zymase and co-enzyme, but they also represent

the concentrations of hexosephosphatase (present in the zymin) and

phosphate (present in the yeast extract), so that at least four

factors were being altered instead of only two.

It has already been mentioned that Euler and Kullberg [1911, 3] found

the conversion of phosphate into hexosephosphate in presence of excess

of glucose to proceed according to a monomolecular reaction (p. 58).

The rate of fermentation is diminished by dilution of the yeast-juice,

but less rapidly than the concentration of the juice. Herzog found

that when the relation between concentration of enzyme and the

velocity constant of the reaction is expressed by the formula

K{1}/K{2} = (C{1}/C{2})^{/n/} where K{1} and K{2} are the velocity

constants corresponding with the enzyme concentrations C{1} and

C{2}, the value for /n/ is 2 for zymin, whilst Euler working with

yeast-juice obtained values varying from 1·29 to 1·67 and decreasing

as K increased.

The temperature coefficient of fermentation by zymin was found [p123]

by Herzog to be K{24·5°}/K{14·5°} = 2·88, which agrees well with the

value found by Slator for yeast-cells (p. 129).

When we endeavour to apply the results of the investigations of the

fermentation of sugar by yeast-juice, zymin, etc., to the process

which goes on in the living cell, considerable difficulties present

themselves. A scheme of fermentation in the living cell can, however,

easily be imagined, which is in harmony with these results. According

to the most simple form of this ideal scheme, the sugar which has

diffused into the cell unites with the fermenting complex and

undergoes the characteristic reaction with phosphate, already present

in the cell, yielding carbon dioxide, alcohol, and hexosephosphate.

The latter is then decomposed, just as it is in yeast-juice, but more

rapidly, and the liberated phosphate again enters into reaction,

partly with the sugar formed from the hexosephosphate and partly

with fresh sugar supplied from outside the cell. The main difference

between fermentation by yeast-juice and by the living cell would then

consist in the rate of decomposition of the hexosephosphate, for it

has been shown that yeast-juice in presence of sufficient phosphate

can ferment sugar at a rate of the same order of magnitude (from 30 to

50 per cent.) as that attained by living yeast.

The difference between the two therefore would appear to lie not so

much in their content of fermenting complex as in their very different

capacity for liberating phosphate from hexosephosphate and thus

supplying the necessary conditions for fermentation.

A simple calculation based on the phosphorus content of living yeast

[Buchner and Haehn, 1910, 2] shows that the whole of this phosphate

must pass through the stage of hexosephosphate every five or six

minutes in order to maintain the normal rate of fermentation, whereas

in an average sample of yeast-juice the cycle, calculated in the same

way, would last nearly two hours.

Wherein this difference resides is a difficult question, which cannot

at present be answered with certainty.

In the first place it must be remembered that a very great

acceleration of the action of the hexosephosphatase is produced

by arsenates (p. 79), and this suggests the possibility that some

substance possessing a similar accelerating power is present in

the yeast-cell and is lost or destroyed in the various processes

involved in rendering the yeast susceptible to phosphate. The great

variety of these processes--extraction of yeast-juice by grinding and

pressing, drying and macerating, heating, treating with acetone and

with toluene--renders this somewhat improbable, and so far no such

substance has been detected. [p124]

A comparison of living yeast, zymin, and yeast-juice shows that these

are situated on an ascending scale with respect to their response to

phosphate. Taking fructose as the substrate in each case, yeast does

not respond to phosphate at all (Slator), the rate of fermentation

by zymin is approximately doubled (p. 46), and that by yeast-juice

increased ten to forty times, whilst the maximum rates are in each

case of the same order of magnitude. Euler and Kullberg, however,

have observed an acceleration of about 25 per cent. in the rate

of fermentation of yeast in presence of a 2 per cent. solution of

monosodium phosphate, NaH{2}PO{4} [1911, 1, 2].

The high rate of fermentation by living yeast and its lack of response

to phosphate may possibly be explained by supposing that the balance

of enzymes in the living cell is such that the supply of phosphate

is maintained at the optimum, and the rate of fermentation cannot

therefore be increased by a further supply.

A further difference lies in the fact that yeast-juice and zymin

respond to phosphate more strongly in presence of fructose than of

glucose, whereas yeast ferments both sugars at the same rate (p. 131),

and this property has been shown to be connected with the specific

relations of fructose to the fermenting complex. It seems possible

that these differences are associated with the gradual passage from

the complete living cell of yeast, through the dead and partially

disorganised cell of zymin to yeast-juice in which the last trace of

cellular organisation has disappeared and the contents of the cell are

uniformly diffused throughout the liquid. Living yeast is, moreover,

not only unaffected by phosphate but only decomposes hexosephosphate

extremely slowly (Iwanoff).

Some light is thrown on these interesting problems by the effect of

antiseptics on fermentation by yeast-cells and by yeast-juice. The

action of toluene has hitherto been most completely studied, and

this substance is an extremely suitable one for the purpose since it

has practically no action whatever on fermentation by yeast-juice.

The experiments of Buchner have, in fact, shown that the normal rate

of fermentation and the total fermentation produced, are almost

unaffected by the presence of toluene even in the proportion of 1 c.c.

to 20 c.c. of yeast-juice. What then is the effect of toluene on the

living yeast-cell? When toluene in large excess is agitated with a

fermenting mixture of yeast and sugar, the rate of fermentation falls

rapidly at first and then more slowly until a relatively constant rate

is attained which gradually decreases in a similar manner to the rate

of fermentation by yeast-juice. Thus at air temperature (16°) 10 grams

of [p125] yeast suspended in 50 c.c. of 6 per cent. glucose solution

gave the following results when agitated with toluene:--

───────────────────────┬───────────────┬───────┬──────────

Time after Addition │ C.c. of CO{2} │ Time. │ C.c. per

of Toluene, Minutes │ per Minute. │ │ Minute

───────────────────────┴───────────────┴───────┴──────────

0 4·6 6 1·6

1 4 8 1·2

2 3·3 12 0·85

3 2·6 24 0·8

4 2 32 0·5

5 1·8 constant

──────────────────────────────────────────────────────────

Simultaneously with this, the yeast acquires the property of

decomposing and fermenting hexosephosphate and of responding to the

addition of phosphate. This last property is only acquired to a small

degree in this way but it becomes much more strongly developed if

the pressed yeast be washed with toluene on the filter pump. Thus 10

grams of yeast after this treatment fermented fructose at 1·2 c.c. per

three minutes; after the addition of phosphate (5 c.c. of 0·6 molar

phosphate) the rate rose to 6·9 and then gradually fell in the typical

manner [Harden, 1910; see also Euler and Johansson, 1912, 3].

The current explanation of the great decrease in rate of fermentation

which attends the action of toluene and other antiseptics on living

yeast, and also follows upon the disintegration of the cell, appears

to be that in living yeast the high rate of fermentation is maintained

by the continued production of relatively large fresh supplies

of fermenting complex, and that when the power of producing this

catalytic agent is destroyed by the poison, the rate of fermentation

falls to a low value, corresponding to the store of zymase still

present in the cell (cf. Buchner, E. and H., and Hahn, 1903, pp. 176,

180).

This explanation implies that the rate of fermentation after the

action of the toluene represents the amount of fermenting complex

present, a supposition which has been shown (p. 53) to be highly

improbable. It further necessitates, as also pointed out independently

by Euler and Ugglas [1911], a rapid destruction of the fermenting

complex both in the process of fermentation and by the action of the

antiseptic, as otherwise the store of zymase remaining in the dead

cell would be practically the same as that contained in the living

cell at the moment when it was subjected to the antiseptic, and this

store would therefore suffice to carry out fermentation at the same

rate in the dead as in the living cell. No such rapid destruction,

however, occurs in yeast-juice, as judged by the rate of fermentation,

which falls off [p126] slowly and to about the same extent in the

presence or absence of toluene. Moreover, as shown above, it is

highly probable that the actual amount of fermenting complex in

yeast-juice is a large fraction of that present at any moment in

the cell, and is capable under suitable conditions of producing

fermentation at a rate comparable with that of the living cell.

This last criticism also applies to the view expressed by Euler

[Euler and Ugglas, 1911; Euler and Kullberg, 1911, 1, 2] that in the

living cell the zymase is partly free and partly combined with the

protoplasm; when the vital activity of the cell is interfered with,

the combined portion of the zymase is thrown out of action and only

that which was free remains active.

The suggestion made by Rubner [1913] that the action of yeast on sugar

is in reality chiefly a vital act, but that a small proportion of

the change is due to enzyme action, is similar in its consequences

to that of Euler and may be met by the same arguments. Buchner and

Skraup [1914] have moreover shown that the effects of sodium chloride

and toluene on the fermenting power of yeast which were observed by

Rubner, can be explained in other ways.

Some other explanation must therefore be sought for this phenomenon.

Great significance must be attached in this connection to the relation

noted above between the degree of disintegration and disorganisation

of the cell and the fall in the normal rate of fermentation. It seems

not impossible that fermentation may be associated in the living cell

with some special structure, or carried on in some special portion

of the cell, perhaps the nuclear vacuole described by Janssens and

Leblanc [1898], Wager [1898, 1911; Wager and Peniston, 1910] and

others which undergoes remarkable changes both during fermentation

and autofermentation [Harden and Rowland, 1901]. The disorganisation

of the cell might lead to many modifications of the conditions,

among others to the dilution of the various catalytic agents by

diffusion throughout the whole volume of the cell. As a matter of

observation the dilution of yeast-juice leads to a considerable

diminution of the rate of fermentation of sugar, and it is possible

that this is one of the chief factors concerned. That phenomena

of this kind may be involved is shown by the remarkable effect of

toluene on the autofermentation of yeast. Whereas the fermentation

of sugar is greatly diminished by the action of toluene, the rate of

autofermentation, which is carried on at the expense of the glycogen

of the cell, is greatly increased. In a typical case, for example, the

autofermentation of 10 grams of yeast suspended in 20 c.c. of water

amounted to 28 c.c. in 4·8 hours [p127] at 25°, whereas the same

amount of yeast in presence of 2 c.c. of toluene gave 97·6 c.c. in

the same time.

Many salts produce a similar effect on English top yeasts (in which

the autofermentation is large) [Harden and Paine, 1912], whereas

Neuberg and Karczag in Berlin [1911, 2] were unable to observe this

phenomenon.

A necessary preliminary of the fermentation of glycogen is its

conversion by a diastatic enzyme into a fermentable sugar, and it

is probable that the effect of the disorganisation of the cell by

toluene is that this enzyme finds more ready access to the glycogen,

which is stored in the plasma of the cell. No such acceleration

of autofermentation is effected by the addition of toluene to

yeast-juice, and hence the result is not due to an acceleration of the

action of the diastatic enzyme on the glycogen.

This effect of toluene is similar in character to the action of

anæsthetics on the leaves of many plants containing glucosides and

enzymes, whereby an immediate decomposition of the glucoside is

initiated [see H. E. and E. F. Armstrong, 1910].

Although as indicated above Euler's theory cannot apply to zymase

itself, if applied to the hexosephosphatase it would afford a

consistent explanation of the facts. According to this modified view

it would be the hexosephosphatase of yeast which existed largely in

the combined form, so that in extracts, in dried yeast and in presence

of toluene only the small fraction which was free would remain active.

The zymase on the other hand would have to be regarded as existing to

a large extent in the free state so that it would pass into extracts

comparatively unimpaired in amount and capable under proper conditions

(i.e. when supplied with sufficient phosphate) of bringing about a

very vigorous fermentation. The theory of combined and free enzymes is

undoubtedly of considerable value, although it cannot be considered as

fully established.

FERMENTATION BY LIVING YEAST.

Much important information as to the nature of the processes involved

in fermentation has been acquired by the direct experimental study of

the action of living yeast on different sugars.

This phenomenon has formed the subject of several investigations

from the kinetic point of view, and its general features may now be

regarded as well established.

The difficulty, which must as far as possible be avoided in

quantitative experiments of this sort with living yeast, is the

alteration [p128] in the amount or properties of the yeast, due to

growth or to some change in the cells. This has been obviated in

the work of Slator [1906] by determining in every case the initial

rate of fermentation, so that the process only continues for a very

short period, during which any change in the amount or constitution

of the yeast is negligible. The method has the further advantage that

interference of the products of the reaction is to a large extent

avoided. The pressure apparatus already described (p. 29) was employed

by Slator, the rate of production of carbon dioxide being measured by

the increase of pressure in the experimental vessel.

INFLUENCE OF CONCENTRATION OF DEXTROSE ON THE RATE OF FERMENTATION.

With regard to this important factor it is found that the action

of living yeast follows the same law as that of most enzymes (p.

121); within certain wide limits the rate of fermentation is almost

independent of the concentration of the sugar. This conclusion has

been drawn by many previous investigators from their experiments

[Dumas, 1874; Tammann, 1889; Adrian Brown, 1892; O'Sullivan, 1898,

1899] and is implicitly contained in the results of Aberson [1903],

although he himself regarded the reaction as monomolecular.

[Illustration: FIG. 8. (Graph Fermentation Velocity vs Conc.

Dextrose--Ed.)]

Slator, working with a suspension of ten to twelve yeast-cells per

1/4000 cubic millimetre at 30°, obtained the results which are

embodied in the curve (Fig. 8).

This shows that, for the amount of yeast in question, the rate of

fermentation is almost constant for concentrations of glucose between

[p129] 1 and 10 grams per 100 c.c., but gradually decreases as the

concentration increases. Below 1 gram per 100 c.c. the rate decreases

very rapidly with the concentration.

It follows from this, in the light of what has already been said (p.

121), that the action of living yeast on sugar follows the same course

as a typical enzyme reaction, although in this case, as in that of

yeast-juice, no information is given as to the exact nature of this

reaction.

INFLUENCE OF THE CONCENTRATION OF YEAST.

It appears to be well established that, when changes in the quantity

and constitution of the yeast employed are eliminated, the rate of

fermentation is exactly proportional to the number of the yeast-cells

present (Aberson, Slator). This result might be anticipated, as

pointed out by Slator, from the fact that the fermentation takes place

within the cell, each cell acting as an independent individual.

The diffusion of sugar into the yeast-cell which necessarily precedes

the act of fermentation has been shown by Slator and Sand [1910] to

occur at such a rate that the supply of sugar is always in excess of

the amount which can be fermented by the cell.

TEMPERATURE COEFFICIENT OF ALCOHOLIC FERMENTATION BY YEAST.

The temperature coefficient of fermentation by living yeast has been

carefully determined by Slator by measurements of the initial rates at

a series of temperatures from 5° to 40° C. The coefficient is found to

be of the same order as that for many chemical reactions, but to vary

considerably with the temperature, a rise in temperature corresponding

with a diminution in the coefficient. The following values were

obtained for glucose; they are independent of the concentration

of yeast and glucose, the class of yeast, and presence or absence

of nutrient salts, and remain the same when inhibiting agents are

present. Almost precisely the same ratios are obtained for fructose

and mannose:--

t. V{/t/ + 5}/V{/t./} V{/t/ + 10}/V{/t./}

5 2·65 5·6

10 2·11 3·8

15 1·80 2·8

20 1·57 2·25

25 1·43 1·95

30 1·35 1·6

35 1·20

Aberson's result, K{/t/ + 10}/K{/t/} = 2·72, which represents the

mean coefficient for 10° between 12° and 33°, agrees well with this.

[p130]

ACTION OF ACCELERATING AGENTS ON LIVING YEAST.

Slator [1908, 1] was unable to find any agent which greatly

accelerated the rate of fermentation of living yeast. Small

concentrations of various inhibiting agents which are often supposed

to act in this way were quite ineffective, and phosphates, which

produce such a striking change in yeast-juice, were almost without

action (cp. p. 124).

Euler and Bäckström [1912], however, have made the important

observation that sodium hexosephosphate causes a considerable

acceleration although it is itself neither fermented nor hydrolysed

under these conditions. The extent of this is evident from the

following numbers:--

────────────────────────────────────────────────────

20 c.c. of 20 per cent. glucose solution.

0·25 g. yeast [Yeast H of St. Erik's brewery].

────────────────────┬───────────────────────────────

Without addition. │ + 0·5 g. Na hexosephosphate.

────────┬───────────┼───────────┬───────────────────

Time. │ CO{2}. │ Time. │ CO{2}.

Min. │ │ Min. │

────────┼───────────┴───────────┴───────────────────

46 │ 10·5 37 8

76 │ 17·5 73 19

197 │ 45 188 52·5

347 │ 74·5 321 123

488 │ 95 450 193·5

────────────────────────────────────────────────────

The observation has been confirmed with English top yeast (Harden and

Young, unpublished experiments), but no explanation of the phenomenon

is at present forthcoming.

Euler has also found [Euler and Cassel, 1913; Euler and Berggren,

1912] that yeast extract, sodium nucleinate and ammonium formate also

increase the rate of fermentation of glucose by yeast, but these

results have been criticised by Harden and Young [1913] on the ground

that the possibility of growth of the yeast during the experiment has

not been excluded.

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:--

┌───────────────────────┬─────────────────────────────────────────┐

│ │ Relative Rates. │

│ Yeast. ├─────────────┬─────────────┬─────────────┤

│ │2·5 per cent.│2·5 per cent.│2·5 per cent.│

│ │ │ │ Glucose + │

│ │ Glucose. │ Mannose. │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].

┌────────────────────┬────────────────────┬───────────────────────┐

│ │ │ Relative Rates. │

│ Yeast. │ Mode of Culture. ├──────────┬────────────┤

│ │ Grown in: │ 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.

┌───────────────────┬────────────────────────────────────┐

│ │ Relative Rates. │

│ Yeast. ├──────────┬────────────┬────────────┤

│ │ 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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Biochem. Zeitsch., 2, 52-80; Zeitsch.

Verein. Rübenzucker-Ind., 1145-1168.

EHRLICH, FELIX (1906, 4), /Zur Frage der Fuselölbildung

der Hefe/ 87

Ber., 39, 4072-4075.

EHRLICH, FELIX (1907, 1), /Ueber die Bedingungen der

Fuselölbildung und über ihren Zusammenhang mit dem

Eiweissaufbau der Hefe/ 87, 94

Ber., 40, 1027-1047; Zeitsch. Verein.

Rübenzucker-Ind., 1907, 461.

EHRLICH, FELIX (1907, 2), /Ueber das natürliche Isomere

des Leucins/ 86

Ber., 40, 2538-2562.

EHRLICH, FELIX (1907, 3), /Die Rolle des Eiweisses und der

Eiweissabbauprodukte bei der Gärung/ 87

Jahrb. d. Versuchs u. Lehranstalt f. Brauerei

in Berlin, 10, 515-529.

EHRLICH, FELIX (1908), /Ueber eine Synthese des Isoleucins/ 86

Ber., 41, 1453-1458; Zeitsch. Verein.

deutsch. Zuckerind., 1908, 528-533.

EHRLICH, FELIX (1909), /Ueber die Entstehung der

Bernsteinsäure bei der alkoholischen Gärung/ 89

Biochem. Zeitsch., 18, 391-423.

EHRLICH, FELIX (1911, 1), /Ueber die Vergärung des

Tyrosins zu p-Oxyphenylethyl alkohol. (Tyrosol)/ 88

Ber., 44, 139-147.

EHRLICH, FELIX (1911, 2), /Ueber die Bildung des

Plasmaeiweisses bei Hefen und Schimmelpilzen/ 89

Biochem. Zeitsch., 36, 477-497.

EHRLICH, FELIX (1912), /Ueber Tryptophol

(β-Indolylæthylalkohol), ein neues Gärprodukt der Hefe aus

Aminosäuren/ 88

Ber., 45, 883-889.

EHRLICH, FELIX, and K. A. JACOBSEN (1911), /Ueber die

Umwandlung von Aminosäuren in Oxysäuren durch Schimmelpilze/ 89

Ber., 44, 888-897.

EHRLICH, FELIX, and P. PISTSCHIMUKA (1912, 1),

/Ueberfuhrung von Aminen in Alkohole durch Hefe und

Schimmelpilze/ 91

Ber., 45, 1006-1012.

EHRLICH, FELIX, and P. PISTSCHIMUKA (1912, 2), /Synthesen

des Tyrosols und seine Umwandlung in Hordenin/ 88

Ber., 45, 2428-2437.

EHRLICH, FELIX (mit A. WENDEL) (1908, 1), /Ueber die

Spaltung racemischer Amidosäuren mittels Hefe, II./ 89

Biochem. Zeitsch., 8, 438-466.

EHRLICH, FELIX, und ADOLPH WENDEL (1908, 2), /Zur Kenntnis

der Leucinfraktion des Eiweisses/ 86

Biochem. Zeitsch., 8, 399-437.

EMMERLING, O. (1899), /Das Verhalten von Glycerinaldehyd

und Dioxyaceton gegen Hefe/ 104

Ber., 32, 542-544.

EMMERLING, O. (1904), /Ueber den Ursprung der Fuselöle/ 86

Ber., 37, 3535-3538.

EMMERLING, O. (1905), /Ueber den Ursprung der Fuselöle/ 86

Ber., 38, 953-956.

EULER, HANS (1905), /Chemische Dynamik der zellfreien

Gärung/ 120

Zeitsch. physiol. Chem., 44, 53-73.

EULER, HANS (1912, 1), /Ueber die Wirkungsweise der

Phosphatese/. (3 Mitteilung) 47, 58

Biochem. Zeitsch., 41, 215-223.

EULER, HANS (1912, 2), /Verhalten der

Kohlenhydratphosphorsäure-ester im Tierkörper. Nach

Versuchen von E. Thorin und D. Johansson/ 51

Zeitsch. physiol. Chem., 79, 375-397.

EULER, HANS (1914), /Ueber die Rolle des Glykogens bei der

Gärung durch lebende Hefe/ 116

Zeitsch. physiol. Chem., 89, 337-344.

EULER, HANS, and BÄCKSTRÖM, HELMA (1912), /Zum Kenntnis

der Hefegärung/. (2 Mitteilung) 47, 68, 130

Zeitsch. physiol. Chem., 77, 394-401.

EULER, HANS, and TH. BERGGREN (1912), /Ueber die primäre

Umwandlung der Hexosen bei der alkoholischen Gärung/ 32, 130

Zeitsch. Gärungsphysiol., 1, 203-218.

EULER, HANS, and HENRY CASSEL (1913), /Ueber Katalysatoren

der alkoholischen Gärung/. Vorläufige Mitteilung 130

Zeitsch. physiol. Chem., 86, 122-129.

EULER, HANS, and A. FODOR (1911), /Ueber ein

Zwischenprodukt der alkoholischen Gärung/ 47, 48, 49, 50, 105

Biochem. Zeitsch., 36, 401-410.

EULER, HANS, and YNGVE FUNKE (1912), /Ueber die Spaltung

der Kohlenhydratphosphorsäure-ester/ 51

Zeitsch. physiol. Chem., 77, 488-496.

EULER, HANS, and DAVID JOHANSSON (1912, 1), /Umwandlung

des Zuckers und Bildung der Kohlensäure bei der

alkoholischen Gärung/ 32

Zeitsch. physiol. Chem., 76, 347-354.

EULER, HANS, and DAVID JOHANSSON (1912, 2),

/Untersuchungen über die chemische Zusammensetzung und

Bildung der Enzyme. IV. Ueber die Anpassung einer Hefe an

Galaktose/ 133

Zeitsch. physiol. Chem., 78, 246-265.

EULER, HANS, and DAVID JOHANNSON (1912, 3), /Ueber den

Einfluss des Toluols auf die Zymase und auf die Phosphatese/ 125

Zeitsch. physiol. Chem., 80, 175-181.

EULER, HANS, and DAVID JOHANSSON (1912, 4), /Versuche über

die enzymatische Phosphatbindung/ 47, 57

Zeitsch. physiol. Chem., 80, 205-211.

EULER, HANS, and DAVID JOHANSSON (1913), /Ueber die

Reaktionsphasen der alkoholischen Gärung/ 47, 52, 54, 73

Zeitsch. physiol. Chem., 85, 192-208.

EULER, HANS, and SIXTEN KULLBERG (1911, 1),

/Untersuchungen über die chemische Zusammensetzung und

Bildung der Enzyme/ 124, 126

Zeitsch. physiol. Chem., 71, 14-30.

EULER, HANS, and SIXTEN KULLBERG (1911, 2), /Ueber das

Verhalten freier und an Protoplasma gebundener

Hefenenzyme/ 124, 126

Zeitsch. physiol. Chem., 73, 85-100 and

partly in Arkiv. Kem. Min. Geol., 4, No. 13, 1-11.

EULER, HANS, and SIXTEN KULLBERG (1911, 3), /Ueber die

Wirkungsweise der Phosphatese/ 47, 58, 122

Zeitsch. physiol. Chem., 74, 15-28.

EULER, HANS, and GUNNAR LUNDEQVIST (1911), /Zur Kenntnis

der Hefegärung/ 131

Zeitsch. physiol. Chem., 72, 97-112.

EULER, HANS, and HJALMAR OHLSÉN (1911), /Ueber den

Einfluss der Temperatur auf die Wirkung der

Phosphatese/ 47, 57, 58

Biochem. Zeitsch., 37, 313-320.

EULER, HANS, and HJALMAR OHLSÉN (1912), /Ueber die

Wirkungsweise der Phosphatese, II/ 47, 57

Zeitsch. physiol. Chem., 76, 468-477.

EULER, HANS, and BETH AF. UGGLAS (1911), /Untersuchungen

über die chemische Zusammensetzung und Bildung der Enzyme./

(2 Mitteilung) 125, 126

Zeitsch. physiol. Chem., 70, 279-290.

FERNBACH, A. (1910), /Sur la dégradation biologique des

hydrates de carbone/ 107

Compt. rend., 151, 1004-1006.

FERNBACH, A., and SCHOEN M. (1913), /L'acide pyruvique,

produit de la vie de la levure/ 109

Compt. rend., 157, 1478-1480.

FISCHER, EMIL, und JULIUS TAFEL (1888), /Oxydation des

Glycerins/ 104

Ber., 21, 2634-2637.

FISCHER, EMIL, und JULIUS TAFEL (1889), /Oxydation des

Glycerins, II./ 104

Ber., 22, 106-110.

FISCHER, EMIL, und HANS THIERFELDER (1894), /Verhalten der

verschiedenen Zucker gegen reine Hefen/ 89

Ber., 27, 2031-2037.

FISCHER, HUGO (1903), /Ueber Enzymwirkung und Gärung/ 19

Centr. Bakt. Par., Abt. II., 10, 547-8.

FITZ, ALB. (1880), /Ueber Spaltpilzgärungen/. (6

Mitteilung) 98

Ber., 13, 1309-1312.

FRANZEN, HARTWIG, and O. STEPPUHN (1911), /Ein Beitrag zur

Kenntnis der alkoholischen Gärung/ 115

Ber., 44, 2915-2919.

FRANZEN, HARTWIG, and O. STEPPUHN (1912, 1),

/Vergärung und Bildung der Ameisensäure durch Hefen/ 103, 115

Zeitsch. physiol. Chem., 77, 129-182.

FRANZEN, HARTWIG, and O. STEPPUHN (1912, 2), /Berichtigung

zu der Abhandlung: Ueber die Vergärung und Bildung der

Ameisensäure durch Hefen/ 115

Zeitsch. physiol. Chem., 78, 164.

GAY-LUSSAC, LOUIS JOSEPH (1810), /Extrait d'un Mémoire sur

la Fermentation/. (Lu à l'Inst., 3 Dec, 1810) 4

Ann. Chim. Phys., 76, 245-259.

GERET, L., und M. HAHN (1898, 1), /Zum Nachweis des im

Hefepresssaft enthaltenen proteolytischen Enzyms/ 20

Ber., 31, 202-205.

GERET, L., und M. HAHN (1898, 2), /Weitere Mitteilungen

über das im Hefepresssaft enthaltene proteolytische Enzym/ 20

Ber., 31, 2335-2344.

GERET, L., und M. HAHN (1900), /Ueber das Hefe-endotrypsin/ 20

Zeitsch. Biologie, 40, 117-172.

GERHARDT, CHARLES (1856), /Traité de Chimie Organique/, 4,

537-546 10, 15

GIGLIOLI, J. (1911), /Della probabile funzione degli olii

essenziali e di altri prodotti volatili delle piante, quale

causa di movemento dei succhi nei tessuli viventi/ 26

Atti. R. Accad. Lincei, 20, II., 349-361.

GREEN, J. REYNOLDS (1897), /The supposed alcoholic enzyme

in yeast/ 19

Annals of Botany, 11, 555-562.

GREEN, J. REYNOLDS (1898), /The alcohol-producing enzyme

of yeast/ 19

Annals of Botany, 12, 491-497.

GROMOFF, T., und O. GRIGORIEFF (1904), /Die Arbeit der

Zymase und der Endotryptase in den abgetöteten Hefezellen

unter verschiedenen Verhältnissen/ 36

Zeitsch. physiol. Chem., 42, 299-329.

GRUBER, M. (1908), /Eduard Buchner/ 18

München, med. Wochensch., 342.

GRÜSS, J. (1904), /Untersuchungen über die Atmung und

Atmungsenzyme der Hefe/ 111

Zeitsch. Ges. Brauwesen, 27, 689.

GRÜSS, J. (1908, 1), /Ueber den Nachweis mittelst

Chromogramm-Methode dass die Hydrogenase aktiv bei der

Alkoholgärung beteiligt ist/ 111

Ber. deutsch, botan. Ges., 26A, 191-196.

GRÜSS, J. (1908, 2), /Hydrogenase oder Reduktase?/ 111

Ber. deutsch, botan. Ges., 26A, 627-630,

Abstract J. Inst. Brewing, 1909, 344.

HAHN, MARTIN (1898), /Das proteolytische Enzym des

Hefepresssaftes/ 20

Ber., 31, 200-201.

HAHN, MARTIN (1908), /Zur Geschichte der Zymaseentdeckung/ 18

München. med. Wochensch., 515.

HANRIOT, M. (1885, 1886), /Sur la décomposition pyrogénée

des acides de la série grasse/ 98

Bull. Soc. Chim., 43, 417; 45, 79-80.

HARDEN, ARTHUR (1901), /The chemical action of Bacillus

coli communis and similar organisms on carbohydrates and

allied compounds/ 90, 115

J. Chem. Soc., 79, 612-628.

HARDEN, ARTHUR (1903), /Ueber alkoholische Gärung mit

Hefepresssaft (Buchner's "Zymase") bei Gegenwart von

Blutserum/ 20, 41

Ber., 36, 715-716.

HARDEN, ARTHUR (1905), /Zymase and alcoholic

fermentation/ 42, 103

J. Inst. Brewing, 11, No. 1.

HARDEN, ARTHUR (1910), /Recent researches on alcoholic

fermentation/ 125

J. Inst. Brewing, 16, 623-639.

HARDEN, ARTHUR (1913), /The enzymes of washed zymin and

dried yeast/ (Lebedeff). /I. Carboxylase/ 82, 83

Biochem. J., 7, 214-217.

HARDEN, ARTHUR, and DOROTHY NORRIS (1912), /The bacterial

production of acetylmethylcarbinol and 2-3 butylene glycol

from various substances/ 110

Proc. Roy. Soc., B., 84, 492-499.

HARDEN, ARTHUR, and ROLAND V. NORRIS (1910), /The

fermentation of galactose by yeast and yeast-juice./

(Preliminary Communication) 32, 133

Proc. Roy. Soc., B., 82, 645-649.

HARDEN, ARTHUR, and ROLAND V. NORRIS (1914), /The enzymes

of washed zymin and dried yeast/ (Lebedeff). /II.

Reductase/ 68, 111, 113

Biochem. J., 8, 100-106.

HARDEN, ARTHUR, and S. G. PAINE (1912), /The action of

dissolved substances on the autofermentation of yeast/ 127

Proc. Roy. Soc., B., 84, 448-459.

HARDEN, ARTHUR, and ROBERT ROBISON (1914), /A new

phosphoric ester obtained by the aid of yeast-juice./

(Preliminary Note) 48

Proc. Chem. Soc., 30, 16-17.

HARDEN, ARTHUR, and SYDNEY ROWLAND (1901),

/Autofermentation and liquefaction of pressed yeast/ 126

J. Chem. Soc., 79, 1227-1235.

HARDEN, ARTHUR, J. THOMPSON, and W. J. YOUNG (1910),

/Apparatus for the collection of gases evolved in

fermentation/ 28

Biochem. J., 5, 230-235.

HARDEN, ARTHUR, and W. J. YOUNG (1902), /Glycogen from

yeast/ 33

J. Chem. Soc., 81, 1224-1233.

HARDEN, ARTHUR, and W. J. YOUNG (1904), /Gärversuche mit

Presssaft aus obergäriger Hefe/ 30, 33, 35

Ber., 37, 1052-1070.

HARDEN, ARTHUR, and W. J. YOUNG (1905, 1), /The alcoholic

ferment of yeast-juice/ 41, 42, 59

J. Physiol., 32. Proceedings of 12 November, 1904.

HARDEN, ARTHUR, and W. J. YOUNG (1905, 2), /The

influence of phosphates on the fermentation of glucose by

yeast-juice./ (Preliminary communication, 1 June, 1905) 42, 47

Proc. Chem. Soc., 21, 189-191.

HARDEN, ARTHUR, and W. J. YOUNG (1906, 1), /The alcoholic

ferment of yeast-juice/ 43

Proc. Roy. Soc., B., 77, 405-420.

HARDEN, ARTHUR, and W. J. YOUNG (1906, 2), /The alcoholic

ferment of yeast-juice. Part II. The co-ferment of

yeast-juice/ 59, 62

Proc. Roy. Soc, B., 78, 369-375.

HARDEN, ARTHUR, and W. J. YOUNG (1906, 3), /Influence

of sodium arsenate on the fermentation of glucose by

yeast-juice/. (Preliminary notice) 37, 75

Proc. Chem. Soc, 22, 283-284.

HARDEN, ARTHUR, and W. J. YOUNG (1907) 65

Biochem. Centr., 6, 888.

HARDEN, ARTHUR, and W. J. YOUNG (1908, 1), /The alcoholic

ferment of yeast-juice. Part III. The function of

phosphates in the fermentation of glucose by

yeast-juice/ 47, 53, 71

Proc. Roy. Soc., B., 80, 299-311.

HARDEN, ARTHUR, and W. J. YOUNG (1908, 2), /The

fermentation of mannose and lævulose by yeast-juice/.

(Preliminary note) 73

Proc. Chem. Soc., 24, 115-117.

HARDEN, ARTHUR, and W. J. YOUNG (1909), /The alcoholic

ferment of yeast-juice. Part IV. The fermentation of

glucose, mannose, and fructose by yeast-juice/ 32, 44, 47, 73

Proc. Roy. Soc., B., 81, 336-347.

HARDEN, ARTHUR, and W. J. YOUNG (1910, 1), /The function

of phosphates in alcoholic fermentation/ 46, 106

Centr. Bakt. Par., Abt. II., 26, 178-184.

HARDEN, ARTHUR, and W. J. YOUNG (1910, 2), /The alcoholic

ferment of yeast-juice. Part V. The function of phosphates

in alcoholic fermentation/ 44, 52, 56

Proc. Roy. Soc., B., 82, 321-330.

HARDEN, ARTHUR, and W. J. YOUNG (1911, 1), /The alcoholic

ferment of yeast-juice. Part VI. The effect of arsenates

and arsenites on the fermentation of the sugars by

yeast-juice/ 56, 75, 77, 79

Proc. Roy. Soc., B., 83, 451-475.

HARDEN, ARTHUR, and W. J. YOUNG (1911, 2), /Ueber die

Zusammensetzung der durch Hefepresssaft gebildeten

Hexosephosphorsäure I/. 47

Biochem. Zeitsch., 32, 173-176.

HARDEN, ARTHUR, and W. J. YOUNG (1912), /Der Mechanismus

der alkoholischen Gärung/ 46, 106, 108

Biochem. Zeitsch., 40, 458-478.

HARDEN, ARTHUR, and W. J. YOUNG (1913), /The enzymatic

formation of polysaccharides by yeast preparations/ 31, 130

Biochem. J., 7, 630-636.

HARDING, VICTOR J. (1912), /The action of enzymes on

hexosephosphate/ 51

Proc. Roy. Soc., B., 85, 418-422.

HELMHOLTZ, HERMANN LUDWIG (1843), /Ueber das Wesen der

Fäulnis und Gärung/ 10

Arch. Anat. Physiol. Joh. Müller, 5, 453-462.

HERZOG, R. O. (1902), /Ueber alkoholische Gärung, I./ 120

Zeitsch. physiol. Chem., 37, 149-160.

HERZOG, R. O. (1904), /Ueber die Geschwindigkeit

enzymatischer Reaktionen/ 120

Zeitsch, physiol. Chem., 41, 416.

HOPPE-SEYLER, F. (1876), /Ueber die Processe der

Gärungen und ihre Beziehungen zum Leben der Organismen/ 14

Pflüger's Archiv, 12, 1-17.

HOPPE-SEYLER, F. (1877), /Ueber Gärungen. Antwort auf

einen Angriff des Herrn Moritz Traube/ 14

Ber., 10, 693-695.

IWANOFF, LEONID (1905), /Ueber Umwandlungen des Phosphors

in der Pflanze/ 47

S. Travaux de la Soc. des Naturalistes de St.

Petersburg, 34.

IWANOFF, LEONID (1907), /Ueber die Synthese der

phospho-organischen Verbindungen in abgetöteten Hefezellen/ 47

Zeitsch. physiol. Chem., 50, 281-288.

IWANOFF, LEONID (1909, 1), /Ueber die Bildung der

phospho-organischen Verbindung und ihre Rolle bei der

Zymasegärung/ 47, 49, 56, 106

Centr. Bakt. Par., Abt. II., 24, 1-12.

IWANOFF, LEONID (1909, 2), /Ueber einen neuen Apparat für

Gärungsversuche/ 29

Centr. Bakt. Par., Abt. II., 24, 429-432.

JANSSENS, A., and A. LEBLANC (1898), /Recherches

cytologiques sous la cellule de levure/ 126

La Cellule, 14, 203-241.

KARAUSCHANOFF, S. (1911), /Zur Frage nach der Bedeutung

des Dioxyacetons als eines intermediären Produktes der

alkoholischen Gärung/ 105

Ber. deutsch, bot. Ges., 29, 322.

KARCZAG, L. (1912, 1), /Ueber die Gärung der verschiedenen

Weinsäuren/ 81

Biochem. Zeitsch., 38, 516-518.

KARCZAG, L. (1912, 2), /In welcher Weise wird die

Weinsäure durch Hefe angegriffen?/ 81

Biochem. Zeitsch., 43, 44-46.

KAYSER, E. (1911), /Sur le suc de levure de bière/ 26

Compt. rend., 152, 1279-1280.

KNOOP, F. (1910), /Ueber den physiologischen Abbau der

Säuren und die Synthese einer Aminosäure im Tierkorper/ 93

Zeitsch. physiol. Chem., 67, 487-502.

KOHL, F. G. (1907), /Ueber das Glykogen und einige

Erscheinungen bei der Sporulation der Hefe/ 116

Ber. deut. bot. Ges., 25, 74-85.

KOHL, F. G. (1909), /Alkoholische Gärung/ 116

Inaug. Diss. Leipzig Abstr. in Zeitsch.

Brauereiwesen, 32, 406-7, and J. Inst.

Brewing, 15, 710-711.

KOSTYTSCHEFF, S. (1912, 1), /Bildung von Acetaldehyd bei

der alkoholischen Zuckergärung/. (Vorläufige Mitteilung) 111

Ber., 45, 1289-1293.

KOSTYTSCHEFF, S. (1912, 2), /Ueber Alkoholgärung/. (1

Mitteilung.) /Ueber die Bildung von Acetaldehyd bei der

alkoholischen Zuckergärung/ 106, 109

Zeitsch. physiol. Chem., 79, 130-145.

KOSTYTSCHEFF, S. (1912, 3), /Ueber Alkoholgärung/. (2

Mitteilung.) /Ueber die Bildung von Äthylalkohol aus

Acetaldehyd durch lebende und getötete Hefe. Von S.

Kostytscheff und E. Hübbenet/ 110

Zeitsch. physiol. Chem., 79, 359-374.

KOSTYTSCHEFF, S. (1913, 1), /Ueber den Mechanismus der

alkoholischen Gärung/ 111

Ber., 46, 339.

KOSTYTSCHEFF, S. (1913, 2), /Ueber Alkoholgärung. III. Die

Bedingungen der Bildung von Acetaldehyd bei der Gärung von

Dauerhefe/ 111, 113

Zeitsch. physiol. Chem., 83, 93-111.

KOSTYTSCHEFF, S. (1914), /Ueber Alkoholgärung/ (6

Mitteilung) /Das Wesen der reduktion von Acetaldehyd durch

lebenden Hefe/ 111

Zeitsch. physiol. Chem., 89, 367-372.

KOSTYTSCHEFF, S., and E. HÜBBENET (1913), /Zur Frage der

Reduktion von Acetaldehyd durch Hefesaft/ 110, 115

Zeitsch. physiol. Chem., 85, 408-411.

KOSTYTSCHEFF, S., and W. BRILLIANT (1913), /Ueber

Alkoholgärung. V. Ueber Eiweissspaltung durch Dauerhefe in

Gegenwart von Zinkchlorid/ 111

Zeitsch. physiol. Chem., 85, 507-516.

KOSTYTSCHEFF, S., and A. SCHELOUMOFF (1913), /Ueber

Alkoholgärung. IV. Ueber Zuckerspaltung durch Dauerhefe in

Gegenwart von Zinkchlorid/ 111, 113

Zeitsch. physiol. Chem., 85, 493-506.

KUNZ, R. (1906), /Ist die bei der alkoholischen Hefegärung

entstehende Bernsteinsäure als Spaltungsprodukt des Zuckers

anzusehen?/ 89

Zeitsch. Unters. Nahr. Genussmittel, 12, 641-645.

KUPFFER, C. V. (1897) 19

München, med. Wochensch., 44, 321-322.

KUSSEROFF, R. (1910), /Eine neue Theorie der alkoholischen

Gärung/ 116

Centr. Bakt. Par., Abt. II., 26, 184-187.

KÜTZING, FRIEDRICH (1837), /Mikroskopische Untersuchungen

über die Hefe und Essigmutter, nebst mehreren anderen

dazugehörigen vegetabilischen Gebilden/. (/Im Auszuge

vorgetragen in der Vers. d. Naturhistor. Vereins des

Harzes, in Alexisbad am 26 Juli, 1837/) 7

J. pr. Chem., 11, 385-409.

LANGE, H. (1898), /Beitrag zur alkoholischen Gärung ohne

Hefezellen/ 19

Wochensch. Brauerei, 15, 377-378.

LAVOISIER, A. (1789), /Traité Élémentaire de Chymie/,

Chap. XIII 2

LEATHES, J. B. (1906), /Problems in Animal Metabolism/ 110

London, 81-86.

LEBEDEFF, ALEX. V. (1908), /Auftreten von Formaldehyd bei

der zellfreien Gärung/ 117

Biochem. Zeitsch., 10, 454-457.

LEBEDEFF, ALEX. V. (1909), /Versuche zur Aufklärung des

zellenfreien Gärungsprocesses mit Hilfe des

Ultrafilters/ 29, 31, 47, 49, 50

Biochem. Zeitsch., 20, 114-125.

LEBEDEFF, ALEX. V. (1910), /Ueber

Hexosephosphorsäureester/. (1 Mitteilung) 31, 47, 48, 50

Biochem. Zeitsch., 28, 213-229.

LEBEDEFF, ALEX. V. (1911, 1), /Ueber den Mechanismus der

alkoholischen Gärung/ 105, 106

Ber., 44, 2932-2942.

LEBEDEFF, ALEX. V. (1911, 2), /Extraction de la zymase par

simple macération/ 24

Compt. rend., 152, 49-51.

LEBEDEFF, ALEX. V. (1911, 3), /Sur l'extraction de la

zymase/ 24

Compt. rend., 152, 1129.

LEBEDEFF, ALEX. V. (1911, 4), /La zymase est-elle une

diastase?/ 34, 35

Ann. Inst. Past., 25, 682-694. (Bull.

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LEBEDEFF, ALEX. V. (1911, 5), /Bemerkungen zu der Arbeit

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WOHL, A., und C. OESTERLIN (1901), /Ueberführung der

Weinsäure in Oxalessigsäure durch Wasserabspaltung bei

niederer Temperatur/ 100

Ber., 34, 1139-1148.

[WÖHLER] (1839), /Das enträtselte Geheimniss der geistigen

Gärung/. (/Vorläufige briefliche Mitteilung./) 8

Annalen, 29, 100-104.

WROBLEWSKI, A. (1898), /Gärung ohne Hefezellen/ 19

Ber., 31, 3218-3225; Centr. Physiol.,

12, 697-701.

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Hefepresssaft/ 19

Centr. Physiol., 13, 284-297.

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(Prel. Note, Proc. Chem. Soc., 1907, 65.) Proc.

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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 ENDNOTE.

Full stops, middle dots "·", or even dots aligned with the top of the

font were variably used in the original as decimal points, and also

for denoting chemical bonding. These have been rendered as middle

dots herein.

Tables and formulas have been edited for clarity and readability,

while honoring the original form. In particular, some tables and

chemical equations were made more compact to control line length.

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.

On page 93, the complex reaction system has been revised into four

simple reactions. The unbalanced original equations have been

retained. Similarly on page 94, the original large diagram was

converted into four simple reactions.

The incorrect formula for the enol II. in the equation for glucose

dehydration near bottom of page 101 was corrected.

Equations (1) on page 108 showing production of glyceric acid from

glyceraldehyde have been simplified and clarified.