Modulation of Phosphoenolpyruvate Synthase Expression Increases
Shikimate Pathway Product Yields in E. coli
Jian Yi, Kai Li, K. M. Draths, and J. W. Frost*
Department of Chemistry, Michigan State University, East Lansing, Michigan 48824
Product yields in microbial synthesis are ultimately limited by the mechanism utilized
for glucose transport. Altered expression of phosphoenolpyruvate synthase was
examined as a method for circumventing these limits. Escherichia coli KL3/pJY1.216A
was cultured under fed-batch fermentor conditions where glucose was the only source
of carbon for the formation of microbial biomass and the synthesis of product
3-dehydroshikimic acid. Shikimate pathway byproducts 3-deoxy-D-arabino-heptulosonic
acid, 3-dehydroquinic acid, and gallic acid were also generated. An optimal expression
level of phosphoenolpyruvate synthase was identified, which did not correspond to
the highest expression levels of this enzyme, where the total yield of 3-dehydroshikimic
acid and shikimate pathway byproducts synthesized from glucose was 51% (mol/mol).
For comparison, the theoretical maximum yield is 43% (mol/mol) for synthesis of
3-dehydroshikimic acid and shikimate pathway byproducts from glucose in lieu of
amplified expression of phosphoenolpyruvate synthase.
An extensive range of chemicals can be synthesized by
exploiting microbial catalysis and the shikimate pathway
(1). Escherichia coli catalyzed syntheses of the amino
acids L-phenylalanine (2) and L-tryptophan (3), as well
as the chiral synthon shikimic acid (4, 5), are already
practiced industrially. Syntheses of commodity chemicals
[adipic acid (6), phenol (7)], pseudocommodity chemicals
[catechol (8), hydroquinone (9), p-hydroxybenzoic acid (10,
11)], fine chemicals [vanillin (12), indigo (13)], and
ultrafine chemicals [gallic acid (14, 15), pyrogallol (14)]
have also been reported. 3-Dehydroshikimic acid (Figure
1) is the most advanced shikimate pathway precursor
common to the synthesis of all of the chemicals cited
above. Strategies elaborated to increase the yield and
concentration of 3-dehydroshikimic acid synthesized from
glucose are thus applicable to the microbial synthesis of
a wide range of molecules.
The mechanism employed to transport carbohydrate
starting material from the culture medium into the
cytoplasm is now recognized to be an important deter-
minant of the yield of a microbially synthesized product
(16-18). In a wide variety of microbes that employ the
phosphoenolpyruvate:carbohydrate phosphotransferase
system (19), one molecule of phosphoenolpyruvate is
converted into pyruvate for each molecule of glucose
transported from the culture medium into the cytoplasm
and phosphorylated to form glucose 6-phosphate (Figure
1). Subsequent conversion of pyruvate to CO
2
results in
a maximum theoretical yield of 43% (mol/mol), based on
a stoichiometric analysis, for the conversion of glucose
into 3-dehydroshikimic acid (20). In this account, the
impact of recycling pyruvate generated by the phospho-
enolpyruvate:carbohydrate phosphotransferase system
back to phosphoenolpyruvate (21, 22) on the yield of
3-dehydroshikimic acid synthesized by E. coli from
glucose is examined under fermentor-controlled condi-
tions. Conversion of phosphotransferase-generated pyru-
* To whom correspondence should be addressed. Tel: (517) 355-
9715 extension 115. Fax: (517) 432-3873. E-mail: frostjw@
argus.cem.msu.edu.
Figure 1. Intermediates (abbreviations): D-glucose 6-phos-
phate (G6P), phosphoenolpyruvate (PEP), D-erythrose 4-phos-
phate (E4P), 3-deoxy-D-arabino-heptulosonic acid (DAH) 7-phos-
phate (DAHP), 3-dehydroquinic acid (DHQ), 3-dehydroshikimic
acid (DHS), gallic acid (GA), shikimic acid (SA). Enzymes
(genes): (a) PEP:carbohydrate phosphotransferase system, (b)
PEP synthase (ppsA), (c) DAHP synthase (aroF
FBR
), (d) DHQ
synthase (aroB), (e) DHQ dehydratase (aroD), (f) shikimate
dehydrogenase (aroE), (g) uncharacterized.
1141 Biotechnol. Prog. 2002, 18, 1141-1148
10.1021/bp020101w CCC: $22.00 © 2002 American Chemical Society and American Institute of Chemical Engineers
Published on Web 10/31/2002