Fermentation Theory
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In biochemistry, fermentation theory refers to the historical study of models of natural fermentation processes, especially
alcoholic Alcoholism is, broadly, any drinking of alcohol that results in significant mental or physical health problems. Because there is disagreement on the definition of the word ''alcoholism'', it is not a recognized diagnostic entity. Predomin ...
and
lactic acid Lactic acid is an organic acid. It has a molecular formula . It is white in the solid state and it is miscible with water. When in the dissolved state, it forms a colorless solution. Production includes both artificial synthesis as well as natu ...
fermentation. Notable contributors to the theory include
Justus Von Liebig Justus Freiherr von Liebig (12 May 1803 – 20 April 1873) was a German scientist who made major contributions to agricultural and biological chemistry, and is considered one of the principal founders of organic chemistry. As a professor at t ...
and
Louis Pasteur Louis Pasteur (, ; 27 December 1822 – 28 September 1895) was a French chemist and microbiologist renowned for his discoveries of the principles of vaccination, microbial fermentation and pasteurization, the latter of which was named afte ...
, the latter of whom developed a purely microbial basis for the fermentation process based on his experiments. Pasteur's work on fermentation later led to his development of the
germ theory of disease The germ theory of disease is the currently accepted scientific theory for many diseases. It states that microorganisms known as pathogens or "germs" can lead to disease. These small organisms, too small to be seen without magnification, invade h ...
, which put the concept of spontaneous generation to rest. Although the fermentation process had been used extensively throughout history prior to the origin of Pasteur's prevailing theories, the underlying biological and chemical processes were not fully understood. In the contemporary, fermentation is used in the production of various alcoholic beverages, foodstuffs, and medications.


Overview of fermentation

Fermentation Fermentation is a metabolic process that produces chemical changes in organic substrates through the action of enzymes. In biochemistry, it is narrowly defined as the extraction of energy from carbohydrates in the absence of oxygen. In food ...
is the anaerobic metabolic process that converts sugar into acids, gases, or alcohols in oxygen starved environments. Yeast and many other microbes commonly use fermentation to carry out
anaerobic respiration Anaerobic respiration is respiration using electron acceptors other than molecular oxygen (O2). Although oxygen is not the final electron acceptor, the process still uses a respiratory electron transport chain. In aerobic organisms undergoing re ...
necessary for survival. Even the human body carries out fermentation processes from time to time, such as during long-distance running; lactic acid will build up in muscles over the course of long-term exertion. Within the human body, lactic acid is the by-product of ATP-producing fermentation, which produces energy so the body can continue to exercise in situations where oxygen intake cannot be processed fast enough. Although fermentation yields less ATP than aerobic respiration, it can occur at a much higher rate. Fermentation has been used by humans consciously since around 5000 BCE, evidenced by jars recovered in the Iran
Zagros Mountains The Zagros Mountains ( ar, جبال زاغروس, translit=Jibal Zaghrus; fa, کوه‌های زاگرس, Kuh hā-ye Zāgros; ku, چیاکانی زاگرۆس, translit=Çiyakani Zagros; Turkish: ''Zagros Dağları''; Luri: ''Kuh hā-ye Zāgro ...
area containing remnants of microbes similar those present in the wine-making process.


History

Prior to Pasteur's research on fermentation, there existed some preliminary competing notions of it. One scientist who had a substantial degree of influence on the theory of fermentation was
Justus von Liebig Justus Freiherr von Liebig (12 May 1803 – 20 April 1873) was a German scientist who made major contributions to agricultural and biological chemistry, and is considered one of the principal founders of organic chemistry. As a professor at t ...
. Liebig believed that fermentation was largely a process of
decomposition Decomposition or rot is the process by which dead organic substances are broken down into simpler organic or inorganic matter such as carbon dioxide, water, simple sugars and mineral salts. The process is a part of the nutrient cycle and is e ...
as a consequence of the exposure of yeast to air and water. This theory was corroborated by Liebig's observation that other decomposing matter, such as rotten plant and animal parts, interacted with sugar in a similar manner as yeast. That is, the decomposition of albuminous matter (i.e. water-soluble proteins) caused sugar to transform to alcohol. Liebig held this view until his death in 1873. A different theory was supported by
Charles Cagniard de la Tour Baron Charles Cagniard de la Tour (31 March 1777 – 5 July 1859) was a French engineer and physicist. Charles Cagniard was born in Paris, and after attending the École Polytechnique became one of the ''ingénieurs géographiques''. He examined t ...
and cell theorist
Theodor Schwann Theodor Schwann (; 7 December 181011 January 1882) was a German physician and physiologist. His most significant contribution to biology is considered to be the extension of cell theory to animals. Other contributions include the discovery of ...
, who claimed that alcoholic fermentation depended on the biological processes carried out by brewer's yeast. Louis Pasteur's interest in fermentation began when he noticed some remarkable properties of
amyl alcohol An amyl alcohol is any of eight alcohols with the formula C5H12O. A mixture of pentyl, amyl alcohols (also called amyl alcohol) can be obtained from fusel alcohol. Amyl alcohol is used as a solvent and in esterification, by which is produced amyl a ...
—a by-product of lactic acid and alcohol fermentation—during his biochemical studies. In particular, Pasteur noted its ability to “rotate the plane of polarized light”, and its “unsymmetric arrangement of atoms." These behaviors were characteristic of organic compounds Pasteur had previously examined, but also presented a hurdle to his own research about a "law of hemihedral correlation". Pasteur had previously been attempting to derive connections between substances' chemical structures and external shape, and the optically active amyl alcohol did not follow his expectations according to the proposed 'law'. Pasteur sought a reason for why there happened to be this exception, and why such a chemical compound was generated during the fermentation process in the first place. In a series of lectures later in 1860, Pasteur attempted to link optical activity and molecular asymmetry to organic origins of substances, asserting that no chemical processes were capable of converting symmetric substances (inorganic) into asymmetric ones (organic). Hence, the amyl alcohol observation provided some of the first motivations for a biological explanation of fermentation. In 1856, Pasteur was able to observe the microbes responsible for alcoholic fermentation under a microscope, as a professor of science in the
University of Lille The University of Lille (french: Université de Lille, abbreviated as ULille, UDL or univ-lille) is a French public research university based in Lille, Hauts-de-France. It has its origins in the University of Douai (1559), and resulted from the m ...
. According to a legend originating in the 1900 biography of Pasteur, one of his chemistry students—an owner of a beetroot alcohol factory in Lille—sought aid from him after an unsuccessful year of brewing. Pasteur performed experiments at the factory in observation of the fermentation process, noticing that yeast globules became elongated after lactic acid was formed, but round and full when alcohol was fermenting correctly. In a different observation, Pasteur inspected particles originating on grapevines under the microscope and revealed the presence of living cells. Leaving these cells immersed in grape juice resulted in active alcoholic fermentation. This observation provided evidence for ending the distinction between ‘artificial’ fermentation in wine and ‘true’ fermentation in yeast products. The previous incorrect distinction had stemmed in part from the fact that yeast had to be added to beer wort in order to provoke desired alcoholic fermentation, while the fermenting catalysts for wine occurred naturally on grapevines; the fermentation of wine had been viewed as 'artificial' since it did not require additional catalyst, but the natural catalyst had been present on the grapevine itself. These observations provided Pasteur with a working hypothesis for future experiments. One of the chemical processes that Pasteur studied was the fermentation of sugar into lactic acid, as occurs in the souring of milk. In an 1857 experiment, Pasteur was able to isolate microorganisms present in lactic acid ferment after the chemical process had taken place. Pasteur then cultivated the microorganisms in a
culture Culture () is an umbrella term which encompasses the social behavior, institutions, and norms found in human societies, as well as the knowledge, beliefs, arts, laws, customs, capabilities, and habits of the individuals in these groups.Tyl ...
with his laboratory. He was then able to accelerate the lactic acid fermentation process in fresh milk by administering the cultivated sample to it. This was an important step in proving his hypothesis that lactic acid fermentation was catalyzed by microorganisms. Pasteur also experimented with the mechanisms of brewer's yeast in the absence of organic nitrogen. By adding pure brewer's yeast to a solution of cane sugar, ammonium salt, and yeast ash, Pasteur was able to observe the alcoholic fermentation process with all of its usual byproducts:
glycerin Glycerol (), also called glycerine in British English and glycerin in American English, is a simple triol compound. It is a colorless, odorless, viscous liquid that is sweet-tasting and non-toxic. The glycerol backbone is found in lipids known ...
,
succinic acid Succinic acid () is a dicarboxylic acid with the chemical formula (CH2)2(CO2H)2. The name derives from Latin ''succinum'', meaning amber. In living organisms, succinic acid takes the form of an anion, succinate, which has multiple biological ro ...
, and small amounts of
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
and fatty matters. However, if any of the ingredients were removed from the solution, no fermentation would occur. To Pasteur, this was proof that yeast required the nitrogen, minerals, and carbon from the medium for its metabolic processes, releasing carbonic acid and ethyl alcohol as byproducts. This also disproved Liebig's theory, since there was no albuminous matter present in the medium; the decomposition of the yeast was not the driving force for the observed fermentation.


Pasteur on spontaneous generation

Before the 1860s and 1870s—when Pasteur published his work on this theory—it was believed that microorganisms and even some small animals such as frogs would spontaneously generate. Spontaneous generation was historically explained in a variety of ways. Aristotle, an ancient Greek philosopher, theorized that creatures appeared out of certain concoctions of earthly elements, such as clay or mud mixing with water and sunlight. Later on, Felix Pouchet argued for the existence of 'plastic forces' within plant and animal debris capable of spontaneously generating eggs, and new organisms were born from these eggs. On top of this, a common piece of evidence that seemed to corroborate the theory was the appearance of maggots on raw meat after it was left exposed to open air. In the 1860s and 1870s, Pasteur's interest in spontaneous generation led him to criticize Pouchet's theories and conduct experiments of his own. In his first experiment, he took boiled sugared yeast-water and sealed it in an airtight contraption. Feeding hot, sterile air into the mixture left it unaltered, while introducing atmospheric dust resulted in microbes and mold appearing within the mixture. This result was also strengthened by the fact that Pasteur used asbestos, a form of totally inorganic matter, to carry the atmospheric dust. In a second experiment, Pasteur used the same flasks and sugar-yeast mixture, but left it idle in 'swan-neck' flasks instead of introducing any extraneous matter. Some flasks were kept open to the common air as the control group, and these exhibited mold and microbial growths within a day or two. When the swan-neck flasks failed to show these same microbial growths, Pasteur concluded that the structure of the necks blocked the passage of atmospheric dust into the solution. From the two experiments, Pasteur concluded that the atmospheric dust carried germs responsible for the 'spontaneous generation' in his broths. Thus, Pasteur's work provided proof that the emergent growth of bacteria in nutrient broths is caused by
biogenesis Spontaneous generation is a superseded scientific theory that held that living creatures could arise from nonliving matter and that such processes were commonplace and regular. It was hypothesized that certain forms, such as fleas, could arise fr ...
rather than some form of spontaneous generation.


Applications

Today, the process of fermentation is used for a multitude of everyday applications including medication, beverages and food. Currently, companies like
Genencor Genencor is a biotechnology company based in Palo Alto, CA and a subsidiary of IFF. Genencor is a producer of Industrial enzymes and low-priced bulk protein. The name Genencor originates with Genencor, Inc., the original joint venture between Gen ...
International uses the production of enzymes involved in fermentation to build a revenue of over $400 million a year. Many medications such as antibiotics are produced by the fermentation process. An example is the important drug
cortisone Cortisone is a pregnene (21-carbon) steroid hormone. It is a naturally-occurring corticosteroid metabolite that is also used as a pharmaceutical prodrug; it is not synthesized in the adrenal glands. Cortisol is converted by the action of the enzy ...
, which can be prepared by the fermentation of a plant steroid known as
diosgenin Diosgenin, a phytosteroid sapogenin, is the product of hydrolysis by acids, strong bases, or enzymes of saponins, extracted from the tubers of ''Dioscorea'' wild yam species, such as the Kokoro. The sugar-free (aglycone) product of such hydrolys ...
. The enzymes used in the reaction are provided by the mold
Rhizopus nigricans ''Rhizopus stolonifer'' is commonly known as white bread mold. It is a member of ''Zygomycota'' and considered the most important species in the genus ''Rhizopus''. It is one of the most common fungi in the world and has a global distribution al ...
. Just as it is commonly known, alcohol of all types and brands are also produced by way of fermentation and
distillation Distillation, or classical distillation, is the process of separation process, separating the components or substances from a liquid mixture by using selective boiling and condensation, usually inside an apparatus known as a still. Dry distilla ...
.
Moonshine Moonshine is high-proof liquor that is usually produced illegally. The name was derived from a tradition of creating the alcohol during the nighttime, thereby avoiding detection. In the first decades of the 21st century, commercial dist ...
is a classic example of how this is carried out. Finally, foods such as yogurt are made by fermentation processes as well. Yogurt is a fermented milk product that contains the characteristic bacterial cultures ''
Lactobacillus bulgaricus ''Lactobacillus delbrueckii'' subsp. ''bulgaricus'' (until 2014 known as ''Lactobacillus bulgaricus'') is one of over 200 published species in the ''Lactobacillus'' genome complex (LGC) and is the main bacterium used for the production of yogurt. ...
'' and ''Streptococcus thermopiles''.


See also

*
Zymotic disease Zymotic disease was a 19th-century medical term for acute infectious diseases, especially "chief fevers and contagious diseases (e.g. typhus and typhoid fevers, smallpox, scarlet fever, measles, erysipelas, cholera, whooping-cough, diphtheria, ...
s (for the
Greek language Greek ( el, label=Modern Greek, Ελληνικά, Elliniká, ; grc, Ἑλληνική, Hellēnikḗ) is an independent branch of the Indo-European family of languages, native to Greece, Cyprus, southern Italy (Calabria and Salento), southern Al ...
term ''zumoun'' for "ferment") *
Cellular respiration Cellular respiration is the process by which biological fuels are oxidised in the presence of an inorganic electron acceptor such as oxygen to produce large amounts of energy, to drive the bulk production of ATP. Cellular respiration may be des ...
*
Fermentation in food processing In food processing, fermentation is the conversion of carbohydrates to alcohol or organic acids using microorganisms—yeasts or bacteria—under anaerobic (oxygen-free) conditions. Fermentation usually implies that the action of microorganism ...
*
Louis Pasteur Louis Pasteur (, ; 27 December 1822 – 28 September 1895) was a French chemist and microbiologist renowned for his discoveries of the principles of vaccination, microbial fermentation and pasteurization, the latter of which was named afte ...
*
Distillation Distillation, or classical distillation, is the process of separation process, separating the components or substances from a liquid mixture by using selective boiling and condensation, usually inside an apparatus known as a still. Dry distilla ...
*
Spontaneous generation Spontaneous generation is a superseded scientific theory that held that living creatures could arise from nonliving matter and that such processes were commonplace and regular. It was hypothesized that certain forms, such as fleas, could arise fr ...


References

{{DEFAULTSORT:Fermentation Theory Obsolete medical theories Microbiology History of science Biology theories Metabolism Louis Pasteur