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Books: The Harvard Classics Volume 38

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We should not be justified in devoting further time to opinions
which are not supported by any serious experiment. Abroad, as
well as in France, the theory of the transformation of albuminous
substances into organized ferments had been advocated long before
it had been taken up by M. Fremy. It no longer commands the
slightest credit, nor do any observers of note any longer give it
the least attention; it might even be said that it has become a
subject of ridicule.

An attempt has also been made to prove that we have contradicted
ourselves, inasmuch as in 1860 we published our opinion that
alcoholic fermentation can never occur without a simultaneous
occurrence of organization, development, and multiplication of
globules; or continued life, carried on from globules already
formed. [Footnote: PASTEUR, Memoire sur la fermentation
alcoolique, 1860: Annales de Chimie et de Physique. The word
globules is here used for cells. In our researches we have always
endeavoured to prevent any confusion of ideas. We stated at the
beginning of our Memoir of 1860 that: "We apply the term
alcoholic to that fermentation which sugar undergoes under the
influence of the ferment known as BEER YEAST." This is, the
fermentation which produces wine and all alcoholic beverages.
This, too, is regarded as the type for a host of similar
phenomena designated, by general usage, under the generic name of
fermentation, and qualified by the name of one of the essential
products of the special phenomenon under observation. Bearing in
mind this fact in reference to the nomenclature that we have
adopted it will be seen that the expression ALCOHOLIC
FERMENTATION cannot be applied to every phenomenon of
fermentation in which alcohol is produced, inasmuch as there may
be a number of phenomena having this character in common. If we
had not at starting defined that particular one amongst the
number of very distinct phenomena, which, to the exclusion of the
others, should bear the name of alcoholic fermentation, we should
inevitably have given rise to a confusion of language that would
soon pass from words to ideas, and tend to introduce unnecessary
complexity into researches which are already, in themselves,
sufficiently complex to necessitate the adoption of scrupulous
care to prevent their becoming still more involved. It seems to
us that any further doubt as to the meaning of the words
ALCOHOLIC FERMENTATION, and the sense in which they are employed,
is impossible, inasmuch as Lavoisier, Gay-Lussac, and Thenard
have applied this term to the fermentation of sugar by means of
beer yeast. It would be both dangerous and unprofitable to
discard the example set by these illustrious masters, to whom we
are indebted for our earliest knowledge of this subject.]
Nothing, however, can be truer than that opinion, and at the
present moment, after fifteen years of study devoted to the
subject since the publication to which we have referred, we need
no longer say, "we think," but instead, "we affirm," that it is
correct. It is, as a matter of fact, to alcoholic fermentation,
properly so called, that the charge to which we have referred
relates--to that fermentation which yields, besides alcohol,
carbonic acid, succinic acid, glycerine, volatile acids, and
other products. This fermentation undoubtedly requires the
presence of yeast--cells under the conditions that we have named.
Those who have contradicted us have fallen into the error of
supposing that the fermentation of fruits is an ordinary
alcoholic fermentation, identical with that produced by beer
yeast, and that, consequently, the cells of that yeast must,
according to own theory, be always present. There is not the
least authority for such a supposition. When we come to exact
quantitative estimations--and these are to be found in the
figures supplied by Messrs. Lechartier and Bellamy--it will be
seen that the proportions of alcohol and carbonic acid gas
produced in the fermentation of fruits differ widely from those
that we find in alcoholic fermentations properly so called, as
must necessarily be the case since in the former the
fermentaction is effected by the cells of a fruit, but in the
latter by cells of ordinary alcoholic ferment. Indeed we have a
strong conviction that each fruit would be found to give rise to
special action, the chemical equation of which would be different
from that in the case of other fruits. As for the circumstance
that the cells of these fruits cause fermentation without
multiplying, this comes under the kind of activity which we have
already distinguished by the expression CONTINUOUS LIFE IN CELLS
ALREADY FORMED.

We will conclude this section with a few remarks on the subject
of equations of fermentations, which have been suggested to us
principally in attempts to explain the results derived from the
fermentation of fruits immersed in carbonic acid gas.

Originally, when fermentations were put amongst the class of
decompositions by contact-action, it seemed probable, and, in
fact, was believed, that every fermentation has its own well-
defined equation which never varied. In the present day, on the
contrary, it must be borne in mind that the equation of a
fermentation varies essentially with the conditions under which
that fermentation is accomplished, and that a statement of this
equation is a problem no less complicated than that in the case
of the nutrition of a living being. To every fermentation may be
assigned an equation in a general sort of way, an equation,
however, which, in numerous points of detail, is liable to the
thousand variations connected with the phenomena of life.
Moreover, there will be as many distinct fermentations brought
about by one ferment as there are fermentable substances capable
of supplying the carbon element of the food of that same ferment,
in the same way that the equation of the nutrition of an animal
will vary with the nature of the food which it consumes. As
regards fermentation producing alcohol, which may be effected by
several different ferments, there will be as in the case of a
given sugar, as many general equations as there are ferments,
whether they be ferment-cells properly so called, or cells of the
organs of living beings functioning as ferments. In the same way
the equation of nutrition varies in the case of different animals
nourished on the same food. And it is from the same reason that
ordinary wort produces such a variety of beers when treated with
the numerous alcoholic ferments which we have described. These
remarks are applicable to all ferments alike; for instance,
butyric ferment is capable of producing a host of distinct
fermentations, in consequence of its ability to derive the
carbonaceous part of its food from very different substances,
from sugar, or lactic acid, or glycerine, or mannite, and many
others.

When we say that every fermentation has its own peculiar ferment,
it must be understood that we are speaking of the fermentation
considered as a whole, including all the accessory products. We
do not mean to imply that the ferment in question is not capable
of acting on some other fermentable substance and giving rise to
fermentation of a very different kind. Moreover, it is quite
erroneous to suppose that the presence of a single one of the
products of a fermentation implies the co-existence of a
particular ferment. If, for example, we find alcohol among the
products of a fermentation, or even alcohol and carbonic acid gas
together, this does not prove that the ferment must be an
alcoholic ferment, belonging to alcoholic fermentations, in the
strict sense of the term. Nor, again, does the mere presence of
lactic acid necessarily imply the presence of lactic ferment. As
a matter of fact, different fermentations may give rise to one or
even several identical products. We could not say with certainty,
from a purely chemical point of view, that we were dealing, for
example, with an alcoholic fermentation properly so called, and
that the yeast of beer must be present in it, if we had not first
determined the presence of all the numerous products of that
particular fermentation under conditions similar to those under
which the fermentation in question had occurred. In works on
fermentation the reader will often find those confusions against
which we are now attempting to guard him. It is precisely in
consequence of not having had their attention drawn to such
observations that some have imagined that the fermentation in
fruits immersed in carbonic acid gas is in contradiction to the
assertion which we originally made in our Memoir on alcoholic
fermentation published in 1860, the exact words of which we may
here repeat:--"The chemical phenomena of fermentation are related
essentially to a vital activity, beginning and ending with the
latter; we believe that alcoholic fermentation never occurs"--we
were discussing the question of ordinary alcoholic fermentation
produced by the yeast of beer--"without the simultaneous
occurrence of organization, development, and multiplication of
globules, or continued life, carried on by means of the globules
already formed. The general results of the present Memoir seem to
us to be it direct opposition to the opinions of MM. Liebig and
Berzelius." These conclusions, we repeat, are as true now as they
ever were, and are as applicable to the fermentation of fruits,
of which nothing was known in 1860, as they are to the
fermentation produced by the means of yeast. Only, in the case of
fruits, it is the cells of the parenchyma that function as
ferment, by a continuation of their activity in carbonic acid gas
whilst in the other case the ferment consists of cells of yeast.

There should be nothing very surprising in the fact that
fermentation can originate in fruits and form alcohol without the
presence of yeast, if the fermentation of fruits were not
confounded completely with alcoholic fermentation yielding the
same products and in the same proportions. It is through the
misuse of words that the fermentation of fruits has been termed
alcoholic, in a way which has misled many persons. [Footnote:
See, for example, the communications of MM. Colin and Poggiale,
and the discussion on them. In the Bulletin de l'Academie de
Medecine, March 2d, 9th, and 30th, and February 16th and 23rd,
1875.] In this fermentation, neither alcohol nor carbonic acid
gas exists in those proportions in which they are found in
fermentation produced by yeast; and, although we may determine in
it the presence of succinic acid, glycerine, and a small quantity
of volatile acids [Footnote: We have elsewhere determined the
formation of minute quantities of volatile acids in alcoholic
fermentation. M. Bechamp, who studied these, recognized several
belonging to the series of fatty acids, acetic acid, butyric acid
&c. "The presence of succinic acid is not accidental, but
constant; if we put aside volatile acids that form in quantities
which we may call infinitely small, we may say that succinic acid
is the only normal acid of alcoholic fermentation."--PASTEUR,
Comptes rendus de l' Academie, t. xlvii., P. 224, 1858] the
relative proportions of these substances will be different from
what they are in the case of alcoholic fermentation.




III. REPLY TO CERTAIN CRITICAL OBSERVATIONS OF THE GERMAN
NATURALISTS, OSCAR BREFELD AND MORITZ TRAUBE.


The essential point of the theory of fermentation which we have
been concerned in proving in the preceding paragraphs may be
briefly put in the statement that ferments properly so called
constitute a class of beings possessing the faculty of living out
of contact with free oxygen; or, more concisely still, we may say
that fermentation is a result of life without air.

If our affirmation were inexact, if ferment cells did require for
their growth or for their increase in number or weight, as all
other vegetable cells do, the presence of oxygen, whether gaseous
or held in solution in liquids, this new theory would lose all
value, its very raison d'etre would be gone, at least as far as
the most important part of fermentations is concerned. This is
precisely what M. Oscar Brefeld has endeavoured to prove in a
Memoir read to the Physico-Medical Society of Wurzburg on July
26th, 1873, in which, although we have ample evidence of the
great experimental skill of its author, he has nevertheless, in
our opinion, arrived at conclusions entirely opposed to fact.

"From the experiments which I have just described," he says, "it
follows, in the most indisputable manner, that A FERMENT CANNOT
INCREASE WITHOUT FREE OXYGEN. Pasteur's supposition that a
ferment, unlike all other living organisms, can live and increase
at the expense of oxygen held in combination, is, consequently,
altogether wanting in any solid basis of experimental proof.
Moreover, since, according to the theory of Pasteur, it is
precisely this faculty of living and increasing at the expense of
the oxygen held in combination that constitutes the phenomenon of
fermentation, it follows that the whole theory, commanding though
it does such general assent, is shown to be untenable; it is
simply inaccurate."

The experiments to which Dr. Brefeld alludes, consisted in
keeping under continued study with the microscope, in a room
specially prepared for the purpose, one or more cells of ferment
in wort in an atmosphere of carbonic acid gas free from the least
traces of free oxygen. We have, however, recognized the fact that
the increase of a ferment out of contact with air is only
possible in the case of a very young specimen; but our author
employed brewer's yeast taken after fermentation, and to this
fact we may attribute the non-success of his growths. Dr.
Brefeld, without knowing it, operated on yeast in one of the
states in which it requires gaseous oxygen to enable it to
germinate again. A perusal of what we have previously written on
the subject of the revival of yeast according to its age will
show how widely the time required for such revival may vary in
different cases. What may be perfectly true of the state of a
yeast to-day may not be so to-morrow, since yeast is continually
undergoing modifications. We have already shown the energy and
activity with which a ferment can vegetate in the presence of
free oxygen, and we have pointed out the great extent to which a
very small quantity; of oxygen held in solution in fermenting
liquids can operate at the beginning of fermentation. It is this
oxygen that produces revival in the cells of the ferment and
enables them to resume the faculty of germinating and continuing
their life, and of multiplying when deprived of air.

In our opinion, a simple reflection should have guarded Dr.
Brefeld against the interpretation which he has attached to his
observations. If a cell of ferment cannot bud or increase without
absorbing oxygen, either free or held in solution in the liquid,
the ratio between the weight of the ferment formed during
fermentation and that of oxygen used up must be constant. We had,
however, clearly established, as far back as 1861, the fact that
this ratio is extremely variable, a fact, moreover, which is
placed beyond doubt by the experiments described in the preceding
section. Though but small quantities of oxygen are absorbed, a
considerable weight of ferment may be generated; whilst if the
ferment has abundance of oxygen at its disposal, it will absorb
much, and the weight of yeast formed will be still greater. The
ratio between the weight of ferment formed and that of sugar
decomposed may pass through all stages within certain very wide
limits, the variations depending on the greater or less
absorption of free oxygen. And in this fact, we believe, lies one
of the most essential supports of the theory which we advocate.
In denouncing the impossibility, as he considered it, of a
ferment living without air or oxygen, and so acting in defiance
of that law which governs all living beings, animal or vegetable,
Dr. Brefeld ought also to have borne in mind the fact which we
have pointed out, that alcoholic yeast is not the only organized
ferment which lives in an anaerobian state. It is really a small
matter that one more ferment should be placed in a list of
exceptions to the generality of living beings, for whom there is
a rigid law in their vital economy which requires for continued
life a continuous respiration, a continuous supply of free
oxygen. Why, for instance, has Dr. Brefeld omitted the facts
bearing on the life of the vibrios of butyric fermentation?
Doubtless he thought we were equally mistaken in these: a few
actual experiments would have put him right.

These remarks on the criticisms of Dr. Brefeld are also
applicable to certain observations of M. Moritz Traube's,
although, as regards the principal object of Dr. Brefeld's
attack, we are indebted to M. Traube for our defence. This
gentleman maintained the exactness of our results before the
Chemical Society of Berlin, proving by fresh experiments that
yeast is able to live and multiply without the intervention of
oxygen. "My researches," he said, "confirm in an indisputable
manner M. Pasteur's assertion that the multiplication of yeast
can take place in media which contain no trace of free oxygen.
... M. Brefeld's assertion to the contrary is erroneous." But
immediately afterwards M. Traube adds: "Have we here a
confirmation of Pasteur's theory? By no means. The results of my
experiments demonstrate on the contrary that this theory has no
true foundation." What were these results? Whilst proving that
yeast could live without air, M. Traube, as we ourselves did,
found that it had great difficulty in living under these
conditions; indeed he never succeeded in obtaining more than the
first stages of true fermentation. This was doubtless for the two
following reasons: first, in consequence of the accidental
production of secondary and diseased fermentations which
frequently prevent the propagation of alcoholic ferment; and,
secondly, in consequence of the original exhausted condition of
the yeast employed. As long ago as 1861, we pointed out the
slowness and difficulty of the vital action of yeast when
deprived of air; and a little way back, in the preceding section,
we have called attention to certain fermentations that cannot be
completed under such conditions without going into the causes of
these peculiarities. M. Traube expresses himself thus: "Pasteur's
conclusion, that yeast in the absence of air is able to derive
the oxygen necessary for its development from sugar, is
erroneous; its increase is arrested even when the greater part of
the sugar still remains undecomposed. IT IS IN A MIXTURE OF
ALBUMINOUS SUBSTANCES THAT YEAST, WHEN DEPRIVED OF AIR, FINDS THE
MATERIALS FOR ITS DEVELOPMENT." This last assertion of M.
Traube's is entirely disproved by those fermentation experiments
in which, after suppressing the presence of albuminous
substances, the action, nevertheless, went on in a purely
inorganic medium, out of contact with air, a fact, of which we
shall give irrefutable proofs. [Footnote: Traube's conceptions
are governed by a theory of fermentation entirely his own, a
hypothetical one, as he admits, of which the following is a brief
summary: "We have no reason to doubt," Traube says, "that the
protoplasm of vegetable cells is itself, or contains within it, a
chemical ferment which causes the alcoholic fermentation of
sugar; its efficacy seems closely connected with the presence of
the cell, inasmuch as, up to the present time, we have discovered
no means of isolating it from the cells with success. In the
presence of air this ferment oxidizes sugar by bringing oxygen to
bear upon it; in the absence of air it decomposes the sugar by
taking away oxygen from one group of atoms of the molecule of
sugar and bringing it to act upon other atoms; on the one hand
yielding a product of alcohol by reduction, on the other hand a
product of carbonic acid gas by oxidation."

Traube supposes that this chemical ferment exists in yeast and in
all sweet fruits, but only when the cells are intact, for he has
proved for himself that thoroughly crushed fruits give rise to no
fermentation whatever in carbonic acid gas. In this respect this
imaginary chemical ferment would differ entirely from those which
we call SOLUBLE FERMENTS, since diastase, emulsine, &c., may be
easily isolated.

For a full account of the views of Brefeld and Traube, and the
discussion which they carried on on the subject of the results of
our experiments, our readers may consult the Journal of the
Chemical Society of Berlin, vii., p. 872. The numbers for
September and December, 1874, in the same volume, contain the
replies of the two authors.]


IV. FERMENTATION OF DEXTRO-TARTRATE OF LIME.

[Footnote: See PASTEUR, Comptes rendus de l'Academie des
Sciences, t. lvi., p. 416.]


Tartrate of lime, in spite of its insolubility in waters is
capable of complete fermentation in a mineral medium.

If we put some pure tartrate of lime, in the form of a
granulated, crystalline powder, into pure water, together with
some sulphate of ammonia and phosphates of potassium and
magnesium, in very small proportions, a spontaneous fermentation
will take place in the deposit in the course of a few days,
although no germs of ferment have been added. A living, organized
ferment, of the vibrionic type, filiform, with tortuous motions,
and often of immense length, forms spontaneously by the
development of some germs derived in some way from the inevitable
particles of dust floating in the air or resting on the surface
of the vessels or material which we employ. The germs of the
vibrios concerned in putrefaction are diffused around us on every
side, and, in all probability, it is one or more of these germs
that develop in the medium in question. In this way they effect
the decomposition of the tartrate, from which they must
necessarily obtain the carbon of their food without which they
cannot exist, while the nitrogen is furnished by the ammonia of
the ammoniacal salt, the mineral principles by the phosphate of
potassium and magnesium, and the sulphur by the sulphate of
ammonia. How strange to see organization, life, and motion
originating under such conditions! Stranger still to think that
this organization, life, and motion are effected without the
participation of free oxygen. Once the germ gets a primary
impulse on its living career by access of oxygen, it goes on
reproducing indefinitely, absolutely without atmospheric air.
Here then we have a fact which it is important to establish
beyond the possibility of doubt, that we may prove that yeast is
not the only organized ferment able to live and multiply when out
of the influence of free oxygen.

Into a flask, like that represented in FIG. 9, of 2.5 litres
(about four pints) in capacity, we put:

Pure, crystallized, neutral tartrate of lime. .. 100 grammes
Phosphate of ammonia. ... . ... . .. ... . ... 1 grammes
Phosphate of magnesium. ... . ... . ... . ... .. 1 grammes
Phosphate of potassium. ... . ... . ... . .. 0.5 grammes
Sulphate of ammonia. ... . ... . ... . ... .. 0.5 grammes
(1 gramme = 15.43 grains)

To this we added pure distilled water, so as entirely to fill the
flask.

In order to expel all the air dissolved in the water and adhering
to the solid substances, we first placed our flask in a bath of
chloride of calcium in a large cylindrical white iron pot set
over a flame. The exit tube of the flask was plunged in a test
tube of Bohemian glass three-quarters full of distilled water,
and also heated by a flame. We boiled the liquids in the flask
and test-tube for a sufficient time to expel all the air
contained in them. We then withdrew the heat from under the test-
tube, and immediately afterwards covered the water which it
contained with a layer of oil and then permitted the whole
apparatus to cool down.

[Illustration with caption: Fig. 9]

Next day we applied a finger to the open extremity of the exit-
tube, which we then plunged in a vessel of mercury. In this
particular experiment which we are describing, we permitted the
flask to remain in this state for a fort-night. It might have
remained there for a century without ever manifesting the least
sign of fermentation, the fermentation of the tartrate being a
consequence of life, and life after boiling no longer existed in
the flask. When it was evident that the contents of the flask
were perfectly inert, we impregnated them rapidly, as follows:
all the liquid contained in the exit-tube was removed by means of
a fine caoutchouc tube, and replaced by about 1 c. (about 17
minims) of liquid and deposit from another flask, similar to the
one we have just described, but which had been fermenting
spontaneously for twelve days; we lost no time in refilling
completely the exit tube with water which had been first boiled
and then cooled down in carbonic acid gas. This operation lasted
only a few minutes. The exit-tube was again plunged under
mercury. Subsequently the tube was not moved from under the
mercury, and as it formed part of the flask, and there was
neither cork nor india-rubber, any introduction of air was
consequently impossible. The small quantity of air introduced
during the impregnation was insignificant and it might even be
shown that it injured rather than assisted the growth of the
organisms, inasmuch as these consisted of adult individuals which
had lived without air and might be liable to be damaged or even
destroyed by it. Be this as it may, in a subsequent experiment we
shall find the possibility removed of any aeration taking place
in this way, however infinitesimal, so that no doubts may linger
on this subject.