Animal infusoria living in the
absence of free oxygen, and the fermentations they bring about
The original French article appears in the Appendix, pages 267-268.
1861, Louis Pasteur
Pasteur, L. 1861. Animalcules infusoires vivant sans gaz oxygene libre
et determinant des fermentations. Comptes rendus de l'Academie des sciences,
Vol. 52, pages 344-347. |
The wide variety of products that are formed during the lactic acid
fermentation are well known. Lactic acid, a gum, mannitol, butyric acid,
alcohol, carbon dioxide and hydrogen all appear simultaneously or successively
in highly variable proportions and in a quite capricious manner. I have
gradually been led to realise that the plant ferment which converts sugar
into lactic acid is different from the one(s) which bring about the production
of the gummy material, and that the latter in turn do not produce lactic
acid. On the other hand it is equally true that none of the various plant
ferments can give rise to butyric acid, if they are separated from the
other forms.
Therefore there has to be a distinct butyric acid ferment. I have been
occupied with this idea for a considerable length of time. I would like
to address myself to the Academy today on the origin of butyric acid in
the so-called lactic acid fermentation.
I will not go into all of the details of this research. I would like
to indicate first one of the conclusions from my work. This is that the
ferment which produces butyric acid is an infusorium.
For a long time I was prevented from discovering this fact because
I had devoted my efforts to eliminating these small animals which I feared
would feed on the plant ferment which I supposed to be the cause of the
butyric acid fermentation and which I was looking for in the liquids which
I was studying. But after unsuccessfully looking for the cause of the butyric
acid fermentation, I was finally struck with the correlation . . . between
this acid and the infusoria, and inversely between the infusoria and the
production of this acid, circumstances which I had attributed to the necessity
of butyric acid for the life of these small animals.
Since then a large number of experiments have convinced me that the
conversion of sugar, mannitol, and lactic acid into butyric acid is due
exclusively to these infusoria, so that it is necessary to consider them
to be the true butyric acid ferment.
Here is their description: They are small cylindrical rods, rounded
at the ends, generally straight, singly or in chains of two, three or four,
and sometimes more. They average about 0.002 mm in diameter, varying in
length from 0.002 to 0.015 or 0.02 mm. They move with a gliding motion.
During this movement, their bodies remain rigid or make slight undulations.
They spin, balancing or quivering actively the two ends of their bodies.
The undulatory nature of their movements becomes very obvious when they
are longer than 0.015 mm in length. Frequently they are bent at one end,
sometimes at both ends. This latter is seldom seen when they are young.
They reproduce by binary fission. It is apparently because of their
mode of reproduction that they occur often in chains. One of the units
attached to others may move quickly several times in order to detach itself....
[These organisms are not animals, as Pasteur supposed, but small motile
bacteria.]
These infusoria can be inoculated in the same way as beer yeast. They
multiply if the medium is suitable for their nutrition. But it should be
stated that they can be inoculated into a medium containing only the crystallizable
and mineral substances sugar, ammonium, and phosphates, and they can reproduce
simultaneously with the rapid appearance of the butyric acid fermentation.
The weight of cells that are formed is significant, although always small
compared to the total quantity of butyric acid formed, but this is true
for all ferments.
The existence of infusoria which are able to bring about fermentations
is already a notable fact. But in addition another unusual aspect should
he mentioned. This is that these infusorial animals are able to live
and multiply
indefinitely in the complete absence of air or free oxygen These infusoria
can not only live in the absence of air, but air actually kills them. If
a stream of carbon dioxide is passed through a medium in which they are
multiplying, their viability and their reproduction are not affected in
the least. On the contrary, under the same conditions, if one substitutes
a stream of air for the carbon dioxide, in one or two hours they all die,
and the butyric acid fermentation which requires their viability is
stopped immediately.
We have arrived therefore at the following, double proposition:
1. The ferment which produces butyric acid is an infusorium.
2. This infusorium lives in the complete absence of free oxygen.
I believe this is the first example known of an animal ferment,
and also
the first example of an animal living in the absence of free oxygen.
We will have to consider how the relationship of the mode of life and
the properties of these small animals, together with the same aspects
of the
plant ferments which can also live without free oxygen, are related
to the processes of fermentation. In the meanwhile I would like to make
no further comment on the ideas which these new facts suggest until further
research has been done.
Commentary by Thomas Brock
This is the first report that any organism can live and reproduce
in the complete absence of free oxygen. cal processes to the presence or
absence of oxygen. This discovery is quite important for general biology,
since it shows that oxygen gas is not a requisite for life. This discovery
opened up for Pasteur a new field of study, relating fermentative and biological
processes to the presence or absence of oxygen. It led to his discovery
that yeast can live either aerobically or anaerobically, and that the yeast
differs in function under these two conditions.
There are a large number of organisms now known that will grow under
anaerobic conditions. Some of these are pathogenic, while others seem to
be saprophytic. So far as is known, oxygen is not toxic to these organisms
in itself, but its presence brings about certain oxidation processes
which allow the accumulation of hydrogen peroxide, which is toxic.
Organisms that can grow either in the presence or absence of free oxygen
are called facultative organisms. Those which will grow only in the absence
of oxygen are called obligate anaerobes. Their culture requires special
procedures that eliminate oxygen gas, and consequently they are not studied
as often as are aerobic and facultative organisms. |