Ethanol fermentation

Ethanol fermentation

Overview

Ethanol fermentation, also referred to as alcoholic fermentation, is a biological process in which sugars such as glucose
Glucose
Glucose is a simple sugar and an important carbohydrate in biology. Cells use it as the primary source of energy and a metabolic intermediate...

, fructose
Fructose
Fructose, or fruit sugar, is a simple monosaccharide found in many plants. It is one of the three dietary monosaccharides, along with glucose and galactose, that are absorbed directly into the bloodstream during digestion. Fructose was discovered by French chemist Augustin-Pierre Dubrunfaut in 1847...

, and sucrose
Sucrose
Sucrose is the organic compound commonly known as table sugar and sometimes called saccharose. A white, odorless, crystalline powder with a sweet taste, it is best known for its role in human nutrition. The molecule is a disaccharide composed of glucose and fructose with the molecular formula...

 are converted into cellular energy and thereby produce ethanol
Ethanol
Ethanol, also called ethyl alcohol, pure alcohol, grain alcohol, or drinking alcohol, is a volatile, flammable, colorless liquid. It is a psychoactive drug and one of the oldest recreational drugs. Best known as the type of alcohol found in alcoholic beverages, it is also used in thermometers, as a...

 and carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

 as metabolic waste products. Because yeast
Yeast
Yeasts are eukaryotic micro-organisms classified in the kingdom Fungi, with 1,500 species currently described estimated to be only 1% of all fungal species. Most reproduce asexually by mitosis, and many do so by an asymmetric division process called budding...

s perform this conversion in the absence of oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...

, ethanol fermentation is classified as anaerobic
Anaerobic respiration
Anaerobic respiration is a form of respiration using electron acceptors other than oxygen. Although oxygen is not used as the final electron acceptor, the process still uses a respiratory electron transport chain; it is respiration without oxygen...

.

Ethanol fermentation occurs in the production of alcoholic beverage
Alcoholic beverage
An alcoholic beverage is a drink containing ethanol, commonly known as alcohol. Alcoholic beverages are divided into three general classes: beers, wines, and spirits. They are legally consumed in most countries, and over 100 countries have laws regulating their production, sale, and consumption...

s and ethanol fuel
Ethanol fuel
Ethanol fuel is ethanol , the same type of alcohol found in alcoholic beverages. It is most often used as a motor fuel, mainly as a biofuel additive for gasoline. World ethanol production for transport fuel tripled between 2000 and 2007 from 17 billion to more than 52 billion litres...

, and in the rising of bread
Bread
Bread is a staple food prepared by cooking a dough of flour and water and often additional ingredients. Doughs are usually baked, but in some cuisines breads are steamed , fried , or baked on an unoiled frying pan . It may be leavened or unleavened...

 dough
Dough
Dough is a paste made out of any cereals or leguminous crops by mixing flour with a small amount of water and/or other liquid. This process is a precursor to making a wide variety of foodstuffs, particularly breads and bread-based items , flatbreads, noodles, pastry, and similar items)...

.

The chemical equation
Chemical equation
A chemical equation is the symbolic representation of a chemical reaction where the reactant entities are given on the left hand side and the product entities on the right hand side. The coefficients next to the symbols and formulae of entities are the absolute values of the stoichiometric numbers...

 below summarizes the fermentation of glucose
Glucose
Glucose is a simple sugar and an important carbohydrate in biology. Cells use it as the primary source of energy and a metabolic intermediate...

, whose chemical formula
Chemical formula
A chemical formula or molecular formula is a way of expressing information about the atoms that constitute a particular chemical compound....

 is C6H12O6.
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Encyclopedia

Ethanol fermentation, also referred to as alcoholic fermentation, is a biological process in which sugars such as glucose
Glucose
Glucose is a simple sugar and an important carbohydrate in biology. Cells use it as the primary source of energy and a metabolic intermediate...

, fructose
Fructose
Fructose, or fruit sugar, is a simple monosaccharide found in many plants. It is one of the three dietary monosaccharides, along with glucose and galactose, that are absorbed directly into the bloodstream during digestion. Fructose was discovered by French chemist Augustin-Pierre Dubrunfaut in 1847...

, and sucrose
Sucrose
Sucrose is the organic compound commonly known as table sugar and sometimes called saccharose. A white, odorless, crystalline powder with a sweet taste, it is best known for its role in human nutrition. The molecule is a disaccharide composed of glucose and fructose with the molecular formula...

 are converted into cellular energy and thereby produce ethanol
Ethanol
Ethanol, also called ethyl alcohol, pure alcohol, grain alcohol, or drinking alcohol, is a volatile, flammable, colorless liquid. It is a psychoactive drug and one of the oldest recreational drugs. Best known as the type of alcohol found in alcoholic beverages, it is also used in thermometers, as a...

 and carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

 as metabolic waste products. Because yeast
Yeast
Yeasts are eukaryotic micro-organisms classified in the kingdom Fungi, with 1,500 species currently described estimated to be only 1% of all fungal species. Most reproduce asexually by mitosis, and many do so by an asymmetric division process called budding...

s perform this conversion in the absence of oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...

, ethanol fermentation is classified as anaerobic
Anaerobic respiration
Anaerobic respiration is a form of respiration using electron acceptors other than oxygen. Although oxygen is not used as the final electron acceptor, the process still uses a respiratory electron transport chain; it is respiration without oxygen...

.

Ethanol fermentation occurs in the production of alcoholic beverage
Alcoholic beverage
An alcoholic beverage is a drink containing ethanol, commonly known as alcohol. Alcoholic beverages are divided into three general classes: beers, wines, and spirits. They are legally consumed in most countries, and over 100 countries have laws regulating their production, sale, and consumption...

s and ethanol fuel
Ethanol fuel
Ethanol fuel is ethanol , the same type of alcohol found in alcoholic beverages. It is most often used as a motor fuel, mainly as a biofuel additive for gasoline. World ethanol production for transport fuel tripled between 2000 and 2007 from 17 billion to more than 52 billion litres...

, and in the rising of bread
Bread
Bread is a staple food prepared by cooking a dough of flour and water and often additional ingredients. Doughs are usually baked, but in some cuisines breads are steamed , fried , or baked on an unoiled frying pan . It may be leavened or unleavened...

 dough
Dough
Dough is a paste made out of any cereals or leguminous crops by mixing flour with a small amount of water and/or other liquid. This process is a precursor to making a wide variety of foodstuffs, particularly breads and bread-based items , flatbreads, noodles, pastry, and similar items)...

.

The chemical process of fermentation of glucose


The chemical equation
Chemical equation
A chemical equation is the symbolic representation of a chemical reaction where the reactant entities are given on the left hand side and the product entities on the right hand side. The coefficients next to the symbols and formulae of entities are the absolute values of the stoichiometric numbers...

 below summarizes the fermentation of glucose
Glucose
Glucose is a simple sugar and an important carbohydrate in biology. Cells use it as the primary source of energy and a metabolic intermediate...

, whose chemical formula
Chemical formula
A chemical formula or molecular formula is a way of expressing information about the atoms that constitute a particular chemical compound....

 is C6H12O6. One mole
Mole (unit)
The mole is a unit of measurement used in chemistry to express amounts of a chemical substance, defined as an amount of a substance that contains as many elementary entities as there are atoms in 12 grams of pure carbon-12 , the isotope of carbon with atomic weight 12. This corresponds to a value...

 of glucose is converted into two moles of ethanol
Ethanol
Ethanol, also called ethyl alcohol, pure alcohol, grain alcohol, or drinking alcohol, is a volatile, flammable, colorless liquid. It is a psychoactive drug and one of the oldest recreational drugs. Best known as the type of alcohol found in alcoholic beverages, it is also used in thermometers, as a...

 and two moles of carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

:
C12H22O11 +H2O + invertase →2 C6H12O6
C6H12O6 + Zymase → 2C2H5OH + 2CO2


C2H5OH is the chemical formula
Chemical formula
A chemical formula or molecular formula is a way of expressing information about the atoms that constitute a particular chemical compound....

 for ethanol
Ethanol
Ethanol, also called ethyl alcohol, pure alcohol, grain alcohol, or drinking alcohol, is a volatile, flammable, colorless liquid. It is a psychoactive drug and one of the oldest recreational drugs. Best known as the type of alcohol found in alcoholic beverages, it is also used in thermometers, as a...

.

Before fermentation takes place, one glucose
Glucose
Glucose is a simple sugar and an important carbohydrate in biology. Cells use it as the primary source of energy and a metabolic intermediate...

 molecule is broken down into two pyruvate molecules. This is known as glycolysis
Glycolysis
Glycolysis is the metabolic pathway that converts glucose C6H12O6, into pyruvate, CH3COCOO− + H+...

. Glycolysis
Glycolysis
Glycolysis is the metabolic pathway that converts glucose C6H12O6, into pyruvate, CH3COCOO− + H+...

 is summarized by the chemical equation
Chemical equation
A chemical equation is the symbolic representation of a chemical reaction where the reactant entities are given on the left hand side and the product entities on the right hand side. The coefficients next to the symbols and formulae of entities are the absolute values of the stoichiometric numbers...

:
C6H12O6 + 2 ADP + 2 Pi + 2 NAD+ → 2 CH3COCOO + 2 ATP + 2 NADH + 2 H2O + 2H+


The chemical formula
Chemical formula
A chemical formula or molecular formula is a way of expressing information about the atoms that constitute a particular chemical compound....

 of pyruvate is CH3COCOO. Pi stands for the inorganic phosphate
Phosphate
A phosphate, an inorganic chemical, is a salt of phosphoric acid. In organic chemistry, a phosphate, or organophosphate, is an ester of phosphoric acid. Organic phosphates are important in biochemistry and biogeochemistry or ecology. Inorganic phosphates are mined to obtain phosphorus for use in...

. As shown by the reaction equation, glycolysis
Glycolysis
Glycolysis is the metabolic pathway that converts glucose C6H12O6, into pyruvate, CH3COCOO− + H+...

 causes the reduction of two molecules of NAD+
Nicotinamide adenine dinucleotide
Nicotinamide adenine dinucleotide, abbreviated NAD, is a coenzyme found in all living cells. The compound is a dinucleotide, since it consists of two nucleotides joined through their phosphate groups. One nucleotide contains an adenine base and the other nicotinamide.In metabolism, NAD is involved...

 to NADH
Nicotinamide adenine dinucleotide
Nicotinamide adenine dinucleotide, abbreviated NAD, is a coenzyme found in all living cells. The compound is a dinucleotide, since it consists of two nucleotides joined through their phosphate groups. One nucleotide contains an adenine base and the other nicotinamide.In metabolism, NAD is involved...

. Two ADP
Adenosine diphosphate
Adenosine diphosphate, abbreviated ADP, is a nucleoside diphosphate. It is an ester of pyrophosphoric acid with the nucleoside adenosine. ADP consists of the pyrophosphate group, the pentose sugar ribose, and the nucleobase adenine....

 molecules are also converted to two ATP
Adenosine triphosphate
Adenosine-5'-triphosphate is a multifunctional nucleoside triphosphate used in cells as a coenzyme. It is often called the "molecular unit of currency" of intracellular energy transfer. ATP transports chemical energy within cells for metabolism...

 and two water molecules via substrate-level phosphorylation
Substrate-level phosphorylation
Substrate-level phosphorylation is a type of metabolism that results in the formation and creation of adenosine triphosphate or guanosine triphosphate by the direct transfer and donation of a phosphoryl group to adenosine diphosphate or guanosine diphosphate from a phosphorylated reactive...

. For more details, refer to the main article on glycolysis
Glycolysis
Glycolysis is the metabolic pathway that converts glucose C6H12O6, into pyruvate, CH3COCOO− + H+...

.

Effect of oxygen


The fermentation process does not require oxygen. If oxygen is present, some species of yeast (Kluyveromyces lactis
Kluyveromyces lactis
Kluyveromyces lactis is a Kluyveromyces yeast commonly used for genetic studies and industrial applications. Its name comes from the ability to assimilate lactose and convert it into lactic acid.- Use :...

, Kluyveromyces lipolytica) oxidize pyruvate completely to carbon dioxide and water. This process is called respiration. Thus these yeasts produce ethanol only in an anaerobic environment.

However, many yeasts such as the commonly used baker's yeast Saccharomyces cerevisiae, and Schizosaccharomyces pombe, prefer fermentation to respiration. These yeasts will produce ethanol even under aerobic conditions given the right sources of nutrition.

Uses



Ethanol fermentation is responsible for the rising of bread dough. Yeast organisms consume sugars in the dough and produce ethanol and carbon dioxide as waste products. The carbon dioxide forms bubbles in the dough, expanding it into something of a foam. Nearly all the ethanol evaporates from the dough when the bread is baked.

All alcoholic beverages, including those produced by carbonic maceration
Carbonic maceration
Carbonic maceration is a winemaking technique, often associated with the French wine region of Beaujolais, in which whole grapes are fermented in a carbon dioxide rich environment prior to crushing. Conventional alcoholic fermentation involves crushing the grapes to free the juice and pulp from the...

, are produced by ethanol fermentation by yeast. Wine and brandy are produced by fermentation of the natural sugars present in fruits, especially grapes. Beer and whiskey are produced by fermentation of grain starches that have been converted to sugar by the enzyme amylase
Amylase
Amylase is an enzyme that catalyses the breakdown of starch into sugars. Amylase is present in human saliva, where it begins the chemical process of digestion. Food that contains much starch but little sugar, such as rice and potato, taste slightly sweet as they are chewed because amylase turns...

, which is present in grain kernels that have been germinated. Amylase-treated grain or amylase-treated potatoes are fermented for the production of vodka. Rum is produced by fermentation of cane sugar. In all cases, the fermentation must take place in a vessel that allows carbon dioxide to escape, but prevents outside air from coming in, as exposure to oxygen would prevent the formation of ethanol.

Similar yeast fermentation of various carbohydrate products is used to produce much of the ethanol used for fuel.

Feedstocks for fuel production


The dominant ethanol feedstock in warmer regions is sugarcane
Sugarcane
Sugarcane refers to any of six to 37 species of tall perennial grasses of the genus Saccharum . Native to the warm temperate to tropical regions of South Asia, they have stout, jointed, fibrous stalks that are rich in sugar, and measure two to six metres tall...

. In temperate regions, sugar beet
Sugar beet
Sugar beet, a cultivated plant of Beta vulgaris, is a plant whose tuber contains a high concentration of sucrose. It is grown commercially for sugar production. Sugar beets and other B...

 is sometimes used instead.

In the United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...

, the main feedstock for the production of ethanol is currently corn. Approximately 2.8 gallons of ethanol are produced from one bushel of corn (0.42 liter per kilogram). While much of the corn turns into ethanol, some of the corn also yields by-products such as DDGS (distillers dried grains with solubles) that can be used as feed for livestock. A bushel of corn produces about 18 pounds of DDGS (320 kilograms of DDGS per metric ton of maize). Although most of the fermentation plants have been built in corn-producing regions, sorghum
Sorghum
Sorghum is a genus of numerous species of grasses, one of which is raised for grain and many of which are used as fodder plants either cultivated or as part of pasture. The plants are cultivated in warmer climates worldwide. Species are native to tropical and subtropical regions of all continents...

 is also an important feedstock for ethanol production in the Plains states. Pearl millet
Pearl millet
Pearl millet is the most widely grown type of millet. Grown in Africa and the Indian subcontinent since prehistoric times, it is generally accepted that pearl millet originated in Africa and was subsequently introduced into India. The center of diversity, and suggested area of domestication, for...

 is showing promise as an ethanol feedstock for the southeastern U.S. and the potential of duckweed is being studied.

In some parts of Europe, particularly France
France
The French Republic , The French Republic , The French Republic , (commonly known as France , is a unitary semi-presidential republic in Western Europe with several overseas territories and islands located on other continents and in the Indian, Pacific, and Atlantic oceans. Metropolitan France...

 and Italy
Italy
Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...

, grapes have become a de facto feedstock for fuel ethanol by the distillation
Distillation
Distillation is a method of separating mixtures based on differences in volatilities of components in a boiling liquid mixture. Distillation is a unit operation, or a physical separation process, and not a chemical reaction....

 of surplus wine
Wine
Wine is an alcoholic beverage, made of fermented fruit juice, usually from grapes. The natural chemical balance of grapes lets them ferment without the addition of sugars, acids, enzymes, or other nutrients. Grape wine is produced by fermenting crushed grapes using various types of yeast. Yeast...

. In Japan
Japan
Japan is an island nation in East Asia. Located in the Pacific Ocean, it lies to the east of the Sea of Japan, China, North Korea, South Korea and Russia, stretching from the Sea of Okhotsk in the north to the East China Sea and Taiwan in the south...

, it has been proposed to use rice normally made into sake as an ethanol source.

Cassava as ethanol feedstock


Ethanol can be made from mineral oil
Mineral oil
A mineral oil is any of various colorless, odorless, light mixtures of alkanes in the C15 to C40 range from a non-vegetable source, particularly a distillate of petroleum....

 or from sugars or starches. Starches are cheapest. The starchy crop with highest energy content per acre is cassava
Cassava
Cassava , also called yuca or manioc, a woody shrub of the Euphorbiaceae native to South America, is extensively cultivated as an annual crop in tropical and subtropical regions for its edible starchy tuberous root, a major source of carbohydrates...

, which grows in tropical countries.

Thailand
Thailand
Thailand , officially the Kingdom of Thailand , formerly known as Siam , is a country located at the centre of the Indochina peninsula and Southeast Asia. It is bordered to the north by Burma and Laos, to the east by Laos and Cambodia, to the south by the Gulf of Thailand and Malaysia, and to the...

 already had a large cassava industry in the 1990s, for use as cattle feed and as a cheap admixture to wheat flour. Nigeria
Nigeria
Nigeria , officially the Federal Republic of Nigeria, is a federal constitutional republic comprising 36 states and its Federal Capital Territory, Abuja. The country is located in West Africa and shares land borders with the Republic of Benin in the west, Chad and Cameroon in the east, and Niger in...

 and Ghana
Ghana
Ghana , officially the Republic of Ghana, is a country located in West Africa. It is bordered by Côte d'Ivoire to the west, Burkina Faso to the north, Togo to the east, and the Gulf of Guinea to the south...

 are already establishing cassava-to-ethanol plants. Production of ethanol from cassava is currently economically feasible when crude oil prices are above US$120 per barrel.

New varieties of cassava are being developed, so the future situation remains uncertain.
Currently, cassava can yield between 25-40 tonnes per hectare (with irrigation and fertilizer), and from a tonne of cassava roots, circa 200 liters of ethanol can be produced (assuming cassava with 22% starch content). A liter of ethanol contains circa 21.46 MJ of energy. The overall energy efficiency of cassava-root to ethanol conversion is circa 32%.

The yeast used for processing cassava is Endomycopsis fibuligera, sometimes used together with bacterium Zymomonas mobilis.

Byproducts of fermentation


Ethanol fermentation produces unharvested byproducts such as heat, food for livestock, and water.

See also

  • Anaerobic respiration
    Anaerobic respiration
    Anaerobic respiration is a form of respiration using electron acceptors other than oxygen. Although oxygen is not used as the final electron acceptor, the process still uses a respiratory electron transport chain; it is respiration without oxygen...

  • Cellular respiration
    Cellular respiration
    Cellular respiration is the set of the metabolic reactions and processes that take place in the cells of organisms to convert biochemical energy from nutrients into adenosine triphosphate , and then release waste products. The reactions involved in respiration are catabolic reactions that involve...

  • Cellulose
    Cellulose
    Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to over ten thousand β linked D-glucose units....

  • Fermentation (biochemistry)
    Fermentation (biochemistry)
    Fermentation is the process of extracting energy from the oxidation of organic compounds, such as carbohydrates, using an endogenous electron acceptor, which is usually an organic compound. In contrast, respiration is where electrons are donated to an exogenous electron acceptor, such as oxygen,...

  • Fermentation (wine)
    Fermentation (wine)
    The process of fermentation in wine turns grape juice into an alcoholic beverage. During fermentation, yeast interact with sugars in the juice to create ethanol, commonly known as ethyl alcohol, and carbon dioxide...