Zymomonas Mobilis
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''Zymomonas mobilis'' is a
Gram negative Gram-negative bacteria are bacteria that, unlike gram-positive bacteria, do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. Their defining characteristic is that their cell envelope consists of ...
, 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: *Adhesive#Anaerobic, Anaerobic ad ...
, non-sporulating, polarly-flagellated, rod-shaped bacterium. It is the only species found in the genus '' Zymomonas''. It has notable
bioethanol Ethanol fuel is fuel containing ethyl alcohol, the same type of alcohol as found in alcoholic beverages. It is most often used as a motor fuel, mainly as a biofuel additive for gasoline. Several common ethanol fuel mixtures are in use a ...
-producing capabilities, which surpass
yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom (biology), kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are est ...
in some aspects. It was originally isolated from alcoholic beverages like the African palm wine, the Mexican
pulque Pulque (; ), occasionally known as octli or agave wine, 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, ...
, 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 Beer is an alcoholic beverage produced by the brewing and fermentation of starches from cereal grain—most commonly malted barley, although wheat, maize (corn), rice, and oats are also used. The grain is mashed to convert starch in the ...
, creating an estery-sulfury flavor due to the production of
acetaldehyde Acetaldehyde (IUPAC systematic name ethanal) is an organic compound, organic chemical compound with the chemical formula, formula , sometimes abbreviated as . It is a colorless liquid or gas, boiling near room temperature. It is one of the most ...
and
hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is toxic, corrosive, and flammable. Trace amounts in ambient atmosphere have a characteristic foul odor of rotten eggs. Swedish chemist ...
. 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 Sucrose, a disaccharide, is a sugar composed of glucose and fructose subunits. It is produced naturally in plants and is the main constituent of white sugar. It has the molecular formula . For human consumption, sucrose is extracted and refined ...
,
glucose Glucose is a sugar with the Chemical formula#Molecular formula, molecular formula , which is often abbreviated as Glc. It is overall the most abundant monosaccharide, a subcategory of carbohydrates. It is mainly made by plants and most algae d ...
, and
fructose Fructose (), or fruit sugar, is a Ketose, ketonic monosaccharide, 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 gal ...
). 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 ...
using the Entner–Doudoroff pathway. The pyruvate is then
fermented Fermentation is a type of anaerobic metabolism which harnesses the redox potential of the reactants to make adenosine triphosphate (ATP) and organic end products. Organic compound, Organic molecules, such as glucose or other sugars, are Catabo ...
to produce
ethanol Ethanol (also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound with the chemical formula . It is an Alcohol (chemistry), alcohol, with its formula also written as , or EtOH, where Et is the ps ...
and
carbon dioxide Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...
as the only products (analogous to yeast). The advantages of ''Z. mobilis'' over '' S. cerevisiae'' with respect to producing
bioethanol Ethanol fuel is fuel containing ethyl alcohol, the same type of alcohol as found in alcoholic beverages. It is most often used as a motor fuel, mainly as a biofuel additive for gasoline. Several common ethanol fuel mixtures are in use a ...
: *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 sugar with the Chemical formula#Molecular formula, molecular formula , which is often abbreviated as Glc. It is overall the most abundant monosaccharide, a subcategory of carbohydrates. It is mainly made by plants and most algae d ...
,
fructose Fructose (), or fruit sugar, is a Ketose, ketonic monosaccharide, 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 gal ...
and
sucrose Sucrose, a disaccharide, is a sugar composed of glucose and fructose subunits. It is produced naturally in plants and is the main constituent of white sugar. It has the molecular formula . For human consumption, sucrose is extracted and refined ...
. Wild-type ''Z. mobilis'' cannot ferment C5 sugars like
xylose Xylose ( , , "wood") is a sugar first isolated from wood, and named for it. Xylose is classified as a monosaccharide of the aldopentose type, which means that it contains five carbon atoms and includes an aldehyde functional group. It is deriv ...
and
arabinose Arabinose is an aldopentose – a monosaccharide containing five carbon atoms, and including an aldehyde (CHO) functional group. Properties For biosynthetic reasons, most saccharides are almost always more abundant in nature as the "D"-form, o ...
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 Acetic acid , systematically named ethanoic acid , is an acidic, colourless liquid and organic compound with the chemical formula (also written as , , or ). Vinegar is at least 4% acetic acid by volume, making acetic acid the main compone ...
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), in the United States 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 Xylitol is a chemical compound with the formula , or HO(CH2)(CHOH)3(CH2)OH; specifically, one particular Stereoisomerism, stereoisomer with that structural formula. It is a colorless or white crystalline solid. It is classified as a polyalcoho ...
, 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. ''Z. mobilis''s
plasma membrane The cell membrane (also known as the plasma membrane or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of a cell from the outside environment (the extr ...
contains
hopanoids Hopanoids are a diverse subclass of Triterpene, triterpenoids with the same hydrocarbon skeleton as the compound hopane. This group of Pentacyclic compound, pentacyclic molecules therefore refers to simple hopenes, hopanols and hopanes, but also to ...
, pentacyclic compounds similar to eukaryotic
sterol A sterol is any organic compound with a Skeletal formula, skeleton closely related to Cholestanol, cholestan-3-ol. The simplest sterol is gonan-3-ol, which has a formula of , and is derived from that of gonane by replacement of a hydrogen atom on ...
s. This allows it to have an extraordinary tolerance to ethanol in its environment, around 13%. ''Z. mobilis'' is traditionally used to make
pulque Pulque (; ), occasionally known as octli or agave wine, 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, ...
.


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 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
{{Authority control Food microbiology Sphingomonadales Ethanol Bacteria described in 1936