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Lactic acid fermentation

From Wikipedia, the free encyclopedia

One isomer of lactic acid

This animation focuses on one molecule of glucose turning into pyruvate then into
lactic acid.In the process there is one 5 carbon glucose molecule and 2 NAD+
molecules. 2 phosphates attach to the ends of the glucose molecule, splitting the
glucose molecule into 2 pyruvate molecules. The NAD+ molecules add another
phosphate onto the open ends of the 2 pyruvate molecules, turning the 2 NAD+ into 2
NADH. Then ADP comes and takes the phosphates, creating 2 ATP molecules.The
pyruvate is turned into 2 lactate molecules.The process then repeats, starting with
another glucose molecule.
Lactic acid fermentation is a metabolic process by which glucose and other six-
carbon sugars (also, disaccharides of six-carbon sugars, e.g. sucrose or lactose)
are converted into cellular energy and the metabolite lactate. It is an anaerobic
fermentation reaction that occurs in some bacteria and animal cells, such as muscle
cells.[1][2][3]

If oxygen is present in the cell, many organisms will bypass fermentation and
undergo cellular respiration; however, facultative anaerobic organisms will both
ferment and undergo respiration in the presence of oxygen.[3] Sometimes even when
oxygen is present and aerobic metabolism is happening in the mitochondria, if
pyruvate is building up faster than it can be metabolized, the fermentation will
happen anyway.

Lactate dehydrogenase catalyzes the interconversion of pyruvate and lactate with


concomitant interconversion of NADH and NAD+.

In homolactic fermentation, one molecule of glucose is ultimately converted to two


molecules of lactic acid. Heterolactic fermentation, in contrast, yields carbon
dioxide and ethanol in addition to lactic acid, in a process called the
phosphoketolase pathway.[1]

Contents
1 History
2 Biochemistry
2.1 Homofermentative process
2.2 Heterofermentative process
3 Applications
3.1 Kimchi
3.2 Sauerkraut
3.3 Sour beer
3.4 Yogurt
3.5 Physiological
4 Notes and references
History
Several chemists discovered during the 19th century some fundamental concepts of
the domain of organic chemistry. One of them for example was the French chemist
Joseph Louis Gay-Lussac, who was especially interested in fermentation processes,
and he passed this fascination to one of his best students, Justus von Liebig. With
a difference of some years, each of them described, together with colleague, the
chemical structure of the lactic acid molecule as we know it today. They had a
purely chemical understanding of the fermentation process, which means that you
can�t see it using a microscope, and that it can only be optimized by chemical
catalyzers. It was then in 1857 when the French chemist Louis Pasteur first
described the lactic acid as the product of a microbial fermentation. During this
time, he worked at the university of Lille, where a local distillery asked him for
advice concerning some fermentation problems. Per chance and with the badly
equipped laboratory he had at that time, he was able to discover that in this
distillery, two fermentations were taking place, a lactic acid one and an alcoholic
one, both induced by some microorganisms. He then continued the research on these
discoveries in Paris, where he also published his theories that presented a stable
contradiction to the purely chemical version represented by Liebig and his
followers. Even though Pasteur described some concepts that are still accepted
nowadays, Liebig refused to accept them until his death in 1873. But even Pasteur
himself wrote that he was �driven� to a completely new understanding of this
chemical phenomenon. Even if Pasteur didn�t find every detail of this process, he
still discovered the main mechanism of how the microbial lactic acid fermentation
works. He was for example the first to describe fermentation as a �form of life
without air."[4][5]

Even if this chemical process hasn�t been properly described before Pasteur�s work,
people had been using microbial lactic acid fermentation for food production much
earlier. Chemical analysis of archeological finds showed that milk fermentation
uses predate the historical period, its first applications were probably a part of
the Neolithic Revolution. Since milk naturally contains lactic acid bacteria, the
discovery of the fermentation process was quite evident, since it happens
spontaneously at an adequate temperature. The problem of these first farmers was
that fresh milk is nearly not digestible by an adult, so they had an interest to
discover this mechanism: In fact, lactic acid bacteria contain the needed enzymes
to digest lactose, and their populations multiply strongly during the fermentation.
Therefore, even short-fermented milk contains enough enzymes to digest the lactose
molecules, once the milk is in the human body, what allowed also adults to consume
it. Even safer was a longer fermentation, which was practiced for cheesemaking.
This process was discovered a very long time ago too, what is proofed by recipes
for cheese production on Cuneiform scripts, the first written documents that exist,
and a bit later in Babylonian and Egyptian texts. What's interesting is the theory
of the competitive advantage of fermented milk products. The idea of this theory is
that the women of these first settled farmer clans could shorten the time between
two children thanks to the additional lactose uptake from milk consumption. This
factor may have given them an important advantage to out-compete the hunter-
gatherer societies.[6]

With the increasing consumption of milk products these societies developed a


lactase persistence by epigenetic inheritance, which means that the milk-digesting
enzyme lactase was present in their body during the whole lifetime, so they could
drink unfermented milk as adults too. This early habituation to lactose consumption
in the first settler societies can still be observed today on regional differences
of this mutation�s concentration. It�s estimated that about 65% of world population
still lacks it.[7] Since these first societies came from regions around eastern
Turkey to central Europe, the gene appears more frequently there and in North
America, as it was settled by Europeans. On the contrary, lactose intolerance is
much more present in Asian countries.

A bottle and glass of Kumis


Milk products and their fermentation have had an important influence on some
cultures� development. This is the case in Mongolia, where people often practice a
pastoral form of agriculture. The milk that they produce and consume in these
cultures is mainly mare milk and has a long tradition. But not every part or
product of the fresh milk has the same meaning. For instance, the fattier part on
the top, the �dee��, is seen as the most valuable part and is therefore often used
to honor guests. Very important with often a traditional meaning as well are
fermentation products of mare milk, like for example the slightly-alcoholic yogurt
kumis. Consumption of these peaks during cultural festivities such as the Mongolian
lunar new year (in spring). The time of this celebration is called the �white
month�, which indicates that milk products (called �white food� together with
starchy vegetables, in comparison to meat products, called �black food�) are a
central part of this tradition. The purpose of these festivities is to �close� the
past year � clean the house or the yurt, honor the animals for having provided
their food, and prepare everything for the coming summer season � to be ready to
�open� the new year. Consuming white food in this festive context is a way to
connect to the past and to a national identity, which is the great Mongolian empire
personified by Genghis Khan. During the time of this empire, the fermented mare
milk was the drink to honor and thank warriors and leading persons, it was not
meant for everybody. Even though it can became a drink for normal people, it has
kept its honorable meaning. Like many other traditions, this one feels the
influence of the globalizing industry. Other products, coming mainly from China and
western countries, like industrial yogurt, tend to replace it more and more, mainly
in urban areas. However, in rural and poorer regions it�s still of great
importance.[8]

Biochemistry
Homofermentative process
Overall, the homofermentative lactic acid fermentation converts a six-carbon sugar
molecule to two lactic acid molecules, storing the released energy into two ATP
molecules. The following equation describes this net result:

{\displaystyle \mathrm {C_{6}H_{12}O_{6}\ {\xrightarrow {2\ ADP\ +\ 2\ P_{i}\


\longrightarrow \ 2\ ATP\ +\ 2\ H_{2}O}}\ 2\ C_{3}H_{6}O_{3}} } {\displaystyle
\mathrm {C_{6}H_{12}O_{6}\ {\xrightarrow {2\ ADP\ +\ 2\ P_{i}\ \longrightarrow \ 2\
ATP\ +\ 2\ H_{2}O}}\ 2\ C_{3}H_{6}O_{3}} }
Heterofermentative process
Heterofermentative bacteria produce one mole of lactate from one mole of glucose as
well as CO2 and acetic acid or ethanol. Ex. Leuconostoc mesenteroides,
Lactobacillus bifermentous, Leconostoc lactis

Applications
Lactic acid fermentation is used in many areas of the world to produce foods that
cannot be produced through other methods.[9][10] The most commercially important
genus of lactic acid-fermenting bacteria is Lactobacillus, though other bacteria
and even yeast are sometimes used.[9] Two of the most common applications of lactic
acid fermentation are in the production of yogurt and sauerkraut.

Pickle

A product prepared by lactic acid bacteria (LAB) fermentation of sugars present in


the pieces of fruits and vegetables. The prepared product is rich in lactic acid,
and only the beneficial bacteria that can tolerate lactic acid pH survive. It not
only assures good quality of nutrients, but it is also a good source of probiotics.

Kimchi
Main article: Kimchi
Kimchi also uses lactic acid fermentation.[11]

Sauerkraut
Main article: Sauerkraut
Lactic acid fermentation is also used in the production of sauerkraut. The main
type of bacteria used in the production of sauerkraut is of the genus Leuconostoc.
[1][12]

As in yogurt, when the acidity rises due to lactic acid-fermenting organisms, many
other pathogenic microorganisms are killed. The bacteria produce lactic acid, as
well as simple alcohols and other hydrocarbons. These may then combine to form
esters, contributing to the unique flavor of sauerkraut.[1]
Sour beer
Main articles: Lambic and Berliner Weisse
Lactic acid is a component in the production of sour beers, including Lambics and
Berliner Weisses.[13]

Yogurt
Main article: Yogurt
The main method of producing yogurt is through the lactic acid fermentation of milk
with harmless bacteria.[9][14] The primary bacteria used are typically
Lactobacillus bulgaricus and Streptococcus thermophilus, and United States as well
as European law requires all yogurts to contain these two cultures (though others
may be added as probiotic cultures).[14] These bacteria produce lactic acid in the
milk culture, decreasing its pH and causing it to congeal. The bacteria also
produce compounds that give yogurt its distinctive flavor. An additional effect of
the lowered pH is the incompatibility of the acidic environment with many other
types of harmful bacteria.[9][14]

For a probiotic yogurt, additional types of bacteria such as Lactobacillus


acidophilus are also added to the culture.[14]

Physiological
Lactobacillus fermentation and accompanying production of acid provides a
protective vaginal microbiome that protects against the proliferation of pathogenic
organisms.[15]

Notes and references


"Fermented Fruits and Vegetables - A Global SO Perspective". United Nations FAO.
1998. Retrieved 2007-06-10.
Ohio State University (1998-04-03). "Glycolysis and Fermentation". Retrieved 2010-
01-12.
Campbell, Neil; Reece, Jane (2005). Biology, 7th Edition. Benjamin Cummings. ISBN
0-8053-7146-X.
Latour, Bruno (1993). Les objets ont-ils une histoire? Rencontre de Pasteur et de
Whitehead dans un bain d�acide lactique. in L�effet Whitehead, Vrin, Paris, pp.196-
217. ISBN 978-2-7116-1216-1.
Benninga, H. (1990). A History of Lactic Acid Making: A Chapter in the History of
Biotechnology, chapter 1 and 2. ISBN 978-0-7923-0625-2.
Shurtleff, William; Aoyagi, Akiko (2004). A Brief History of Fermentation, East
and West. In History of Soybeans and Soyfoods, 1100 B.C. to the 1980s. ISBN 1-
58008336-6.
Br�ssow, Harald (2013). Nutrition, population growth and disease: a short history
of lactose. in Environmental Microbiology Volume 15, pages 2154-2161.
Ruhlmann, Sandrine; Gardelle, Linda (2013). Les dessus et les dessous du lait.
Sociologie et politique du lait et de ses d�riv�s en Mongolie. in �tudes mongoles
et sib�riennes, centrasiatiques et tib�taines, n� 43-44.
"Lactic acid fermentation". Retrieved 2010-01-09.
"Lactic acid fermentation". Archived from the original on 2009-08-02. Retrieved
2010-01-09.
"Lactic acid fermentation in the production of foods from vegetables, cereals and
legumes". Antonie van Leeuwenhoek. Antonie van Leeuwenhoek Journal. 49 (3):
337�348. 1983. doi:10.1007/BF00399508. Retrieved 2010-06-28.
"Sauerkraut Fermentation". University of Wisconsin�Madison. 1999. Retrieved 2010-
01-09.
Nummer, Brian A. "Brewing With Lactic Acid Bacteria". MoreFlavor Inc. Retrieved 2
October 2013.
"Yogurt Production". 2006-12-29. Retrieved 2010-01-09.
Nardis, C.; Mastromarino, P.; Mosca, L. (September�October 2013). "Vaginal
microbiota and viral sexually transmitted diseases". Annali di Igiene. 25 (5):
443�56. doi:10.7416/ai.2013.1946. PMID 24048183.
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Categories: FermentationMetabolic pathways
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