Xylose metabolism
D-Xylose is a five-carbon aldose (pentose, monosaccharide) that can be catabolized or metabolized into useful products by a variety of organisms.
There are at least four different pathways for the catabolism of D-xylose: An oxido-reductase pathway is present in eukaryotic microorganisms. Prokaryotes typically use an isomerase pathway, and two oxidative pathways, called Weimberg and Dahms pathways respectively, are also present in prokaryotic microorganisms.
Pathways
The oxido-reductase pathway
This pathway is also called the “Xylose Reductase-Xylitol Dehydrogenase” or XR-XDH pathway.
The isomerase pathway
In this pathway the enzyme
Weimberg pathway
The Weimberg pathway[2] is an oxidative pathway where the D-xylose is oxidized to D-xylono-lactone by a D-xylose dehydrogenase followed by a lactonase to hydrolyze the lactone to D-xylonic acid. A xylonate dehydratase is splitting off a water molecule resulting in 2-keto 3-deoxy-xylonate. 2-keto-3-deox-D-xylonate dehydratase forms the α-ketoglutarate semialdehyde. This is subsequently oxidised via α-ketoglutarate semialdehyde dehydrogenase to yield 2-ketoglutarate which serves as a key intermediate in the citric acid cycle.[3]
Dahms pathway
The Dahms pathway
Biotechnological applications
It is desirable to ferment D-xylose to ethanol. This can be accomplished either by native xylose fermenting yeasts such as Scheffersomyces
In another approach, bacterial xylose isomerases have been introduced into S. cerevisiae. This enzyme catalyze the direct formation of D-xylulose from D-xylose. Many attempts at expressing bacterial isomerases were not successful due to misfolding or other problems, but a xylose isomerase from the anaerobic fungus Piromyces Sp. has proven effective.[6] One advantage claimed for S. cerevisiae engineered with the xylose isomerase is that the resulting cells can grow anaerobically on xylose after evolutionary adaptation.
Studies on
Since the pentose phosphate pathway produces additional NADPH during metabolism, limiting this step will help to correct the already evident imbalance between NAD(P)H and NAD+ cofactors and reduce xylitol byproduct formation.Another experiment comparing the two D-xylose metabolizing pathways revealed that the XI pathway was best able to metabolize D-xylose to produce the greatest ethanol yield, while the XR-XDH pathway reached a much faster rate of ethanol production.[8]
Overexpression of the four genes encoding non-oxidative
The aim of this genetic recombination in the laboratory is to develop a yeast strain that efficiently produces ethanol. However, the effectiveness of D-xylose metabolizing laboratory strains do not always reflect their metabolism abilities on raw xylose products in nature. Since D-xylose is mostly isolated from agricultural residues such as wood stocks then the native or genetically altered yeasts will need to be effective at metabolizing these less pure natural sources.
Varying expression of the XR and XDH enzyme levels have been tested in the laboratory in the attempt to optimize the efficiency of the D-xylose metabolism pathway.[12]