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''Zymomonas mobilis'' is a
Gram negative The gram (originally gramme; SI unit symbol g) is a unit of mass in the International System of Units (SI) equal to one one thousandth of a kilogram. Originally defined as of 1795 as "the absolute weight of a volume of pure water equal to th ...
, facultative
anaerobic Anaerobic means "living, active, occurring, or existing in the absence of free oxygen", as opposed to aerobic which means "living, active, or occurring only in the presence of oxygen." Anaerobic may also refer to: * Anaerobic adhesive, a bonding a ...
, non-sporulating, polarly-flagellated, rod-shaped bacterium. It is the only species found in the genus '' Zymomonas''. It has notable
bioethanol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a h ...
-producing capabilities, which surpass
yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constitut ...
in some aspects. It was originally isolated from alcoholic beverages like the African palm wine, the Mexican
pulque Pulque (; nci, metoctli), or octli, is an alcoholic beverage made from the fermented sap of the maguey (agave) plant. It is traditional in central Mexico, where it has been produced for millennia. It has the color of milk, a rather viscous co ...
, and also as a contaminant of cider and beer (cider sickness and beer spoilage) in European countries.


Beer spoilage

''Zymomonas'' is an unwanted waterborn bacteria in beer, creating an estery-sulfury flavor due to the production of
acetaldehyde Acetaldehyde (IUPAC systematic name ethanal) is an organic chemical compound with the formula CH3 CHO, sometimes abbreviated by chemists as MeCHO (Me = methyl). It is a colorless liquid or gas, boiling near room temperature. It is one of the m ...
and hydrogen sulfide. This can be likened to a rotten apple smell or fruity odor. ''Zymomonas'' have not been reported in lager breweries due to the low temperatures (8–12 °C) and stringent carbohydrate requirements (able to ferment only sucrose,
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, using ...
, and
fructose Fructose, or fruit sugar, is a ketonic simple sugar found in many plants, where it is often bonded to glucose to form the disaccharide sucrose. It is one of the three dietary monosaccharides, along with glucose and galactose, that are absorbe ...
). It is commonly found in cask-conditioned ales where priming sugar is used to carbonate the beer. The optimum growth temperature is 25 to 30 °C.


Ethanol production

''Zymomonas mobilis'' degrades sugars to
pyruvate Pyruvic acid (CH3COCOOH) is the simplest of the alpha-keto acids, with a carboxylic acid and a ketone functional group. Pyruvate, the conjugate base, CH3COCOO−, is an intermediate in several metabolic pathways throughout the cell. Pyruvic ac ...
using the
Entner–Doudoroff pathway The Entner–Doudoroff pathway (ED Pathway) is a metabolic pathway that is most notable in Gram-negative bacteria, certain Gram-positive bacteria and archaea. Glucose is the substrate in the ED pathway and through a series of enzyme assisted che ...
. The pyruvate is then
fermented 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 p ...
to produce
ethanol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a hyd ...
and carbon dioxide as the only products (analogous to yeast). The advantages of ''Z. mobilis'' over '' S. cerevisiae'' with respect to producing
bioethanol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a h ...
: *higher sugar uptake and ethanol yield (up to 2.5 times higher), *lower biomass production, *higher ethanol tolerance up to 16% (v/v), *does not require controlled addition of oxygen during the fermentation, However, in spite of these attractive advantages, several factors prevent the commercial usage of ''Z. mobilis'' in
cellulosic ethanol Cellulosic ethanol is ethanol (ethyl alcohol) produced from cellulose (the stringy fiber of a plant) rather than from the plant's seeds or fruit. It can be produced from grasses, wood, algae, or other plants. It is generally discussed for use as a ...
production. The foremost hurdle is that its substrate range is limited to
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, using ...
,
fructose Fructose, or fruit sugar, is a ketonic simple sugar found in many plants, where it is often bonded to glucose to form the disaccharide sucrose. It is one of the three dietary monosaccharides, along with glucose and galactose, that are absorbe ...
and sucrose. Wild-type ''Z. mobilis'' cannot ferment C5 sugars like xylose and arabinose which are important components of lignocellulosic hydrolysates. Unlike ''E. coli'' and yeast, ''Z. mobilis'' cannot tolerate toxic inhibitors present in lignocellulosic hydrolysates such as acetic acid and various phenolic compounds. Concentration of acetic acid in lignocellulosic hydrolysates can be as high as 1.5% (w/v), which is well above the tolerance threshold of ''Z. mobilis''. Several attempts have been made to engineer ''Z. mobilis'' to overcome its inherent deficiencies. National Renewable Energy Laboratory (NREL), USA has made significant contributions in expanding its substrate range to include C5 sugars like xylose and arabinose. Acetic acid resistant strains of ''Z. mobilis'' have been developed by rational metabolic engineering efforts, mutagenesis techniques or adaptive mutation. However, when these engineered strains metabolize mixed sugars in presence of inhibitors, the yield and productivity are much lower, thus preventing their industrial application. An extensive adaptation process was used to improve xylose fermentation in ''Z. mobilis''. By adapting a strain in a high concentration of xylose, significant alterations of metabolism occurred. One noticeable change was reduced levels of xylitol, a byproduct of xylose fermentation which can inhibit the strain’s xylose metabolism. One of the reasons for lower xylitol production was mutation in a putative gene encoding for an aldo-keto reductase that catalyzes the reduction of xylose to xylitol. An interesting characteristic of ''Z. mobilis'' is that its
plasma membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment (the ...
contains
hopanoids Hopanoids are a diverse subclass of triterpenoids with the same hydrocarbon skeleton as the compound hopane. This group of pentacyclic molecules therefore refers to simple hopenes, hopanols and hopanes, but also to extensively functionalized deriva ...
, pentacyclic compounds similar to eukaryotic
sterol Sterol is an organic compound with formula , whose molecule is derived from that of gonane by replacement of a hydrogen atom in position 3 by a hydroxyl group. It is therefore an alcohol of gonane. More generally, any compounds that contain the gon ...
s. This allows it to have an extraordinary tolerance to ethanol in its environment, around 13%. Zymomonas mobilis is traditionally used to make
pulque Pulque (; nci, metoctli), or octli, is an alcoholic beverage made from the fermented sap of the maguey (agave) plant. It is traditional in central Mexico, where it has been produced for millennia. It has the color of milk, a rather viscous co ...
.


Genome

The genome of ''Z. mobilis'' strain ZM4 has been sequenced and contains 2,056,416 bp encoding 1,998 protein coding genes. This revealed that ''Z. mobilis'' can only metabolise glucose via the
Entner–Doudoroff pathway The Entner–Doudoroff pathway (ED Pathway) is a metabolic pathway that is most notable in Gram-negative bacteria, certain Gram-positive bacteria and archaea. Glucose is the substrate in the ED pathway and through a series of enzyme assisted che ...
and is not capable of using the Embden–Meyerhof–Parnas pathway.


References


External links


Ethanol fermentation technology—''Zymomonas mobilis''''Zymomonas mobilis'' ZM4 genome pageType strain of ''Zymomonas mobilis'' at Bac''Dive'' - the Bacterial Diversity Metadatabase
{{Taxonbar, from=Q245744 Sphingomonadales Ethanol Bacteria described in 1936