Increased Phenylalanine Production by Growing and Nongrowing
Escherichia coli Strain CWML2
Christian Weikert, Uwe Sauer, and James E. Bailey*
Institute of Biotechnology, ETH Zu ¨ rich, CH-8093 Zu ¨ rich, Switzerland
Chemostat selection at low dilution rate in glycerol-limited minimal medium was
previously employed to isolate the mutant Escherichia coli strain CWML2 which
exhibits shorter lag phases, decreased acetate production, and higher specific growth
rates and biomass yields in batch culture (Weikert, C.; Sauer, U.; Bailey, J. E.
Microbiology 1997, 143, 1567-1574). In this study, CWML2 was analyzed for its
biochemical production capabilities in batch culture and under nongrowing conditions.
Both CWML2 and MG1655 were transformed with plasmid pSY130-14, which encodes
feedback resistant mutants of the enzymes chorismate mutase P-prephenate synthase
and 3-hydroxy-D-arabinoheptulosonate-7-phosphate dehydratase, to enable phenyla-
lanine production. In batch culture, transformed CWML2 produced twice as much
phenylalanine as did MG1655:pSY130-14. In contrast to the reference strain,
substantial growth-independent production of phenylalanine was calculated for
CWML2:pSY130-14 by using Luedeking-Piret kinetic analysis. Over a period of 30
h, nongrowing cells of CWML2:pSY130-14 exhibited a 2.5-fold higher specific pheny-
lalanine production rate. The apparent capability of E. coli CWML2 to partly uncouple
metabolic activity from growth suggests a potentially general advantage of this class
of modified hosts for production of biochemicals.
Introduction
In nature, microorganisms are usually faced with
severe and persistent nutrient shortage interrupted by
short periods of abundant nutrient supply (Gottschal,
1992). As an adaptation to these conditions, most
microorganisms have developed strategies for survival
in a metabolically inert state (Chesbro et al., 1990;
Jenkins et al., 1988; Matin, 1992). Upon encountering
substrate availability, they resume growth at the maxi-
mum possible rates, often accompained by extensive
overflow metabolism and reduced bioenergetic efficiency
(Russel and Cook, 1995). For many biotechnological
applications, however, desirable process organisms should
exhibit high metabolic activity under slow-growing or
nongrowing conditions and yet maintain tight control of
overflow metabolism. Therefore, control of substrate
uptake and catabolism in the absence of growth is a key
factor for improving the productivity and operational life
span of microbial biocatalysts (Matin, 1992).
Several strategies for improving productivity by reduc-
ing biomass formation have been elucidated, including
the utilization of slow-growing, immobilized (Bailey et
al., 1987; Karkare et al., 1986), or stationary-phase cells
(Tunner et al., 1992). Upon entry into stationary phase
the cells undergo physiological changes that confer
resistance against adverse physiological conditions (Ma-
tin, 1991). Unfortunately these changes are associated
with a reduction in their specific biocatalytic activity, as
was shown in slow-growing chemostat culture (Flickinger
and Rouse, 1993). Complete separation of biomass and
product formation, however, would dramatically improve
the commercial viability of many microbial processes, if
suitable organisms could be identified or genetically
engineered (Matin, 1991).
Recently, we have been able to isolate Escherichia coli
strains that exhibited generally improved physiological
properties (Weikert et al., 1997). These strains were
isolates from the dominant population of a glycerol-
limited, long-term chemostat after 219 generations that
was operated at an extremly low dilution rate of 0.05 h
-1
.
Proteome analysis using two-dimensional gel electro-
phoresis revealed a very high degree of similarity be-
tween the parental MG1655 strain and the chemostat
isolates, thus proving the close relation of the isolated
strains to their progenitor. One of the selected strains,
E. coli CWML2, showed higher specific growth rates and
increased biomass yields on glycerol, glucose, and the
gluconeogenic substrate acetate (Weikert et al., 1997).
This phenotypic property was combined with shorter lag
phases and increased resistance to various stresses.
Furthermore, this strain was shown to produce signifi-
cantly higher concentrations of a heterologous secreted
protein than the progenitor after transformation of both
with the same expression vector (Weikert et al., 1998).
In this paper, we examine the extent to which the
improved physiological properties of E. coli strain CW-
ML2 may be exploited for production of primary meta-
bolites. The progenitor, wild-type E. coli MG1655, was
used as the reference organism. As an example of a
biotechnologically relevant metabolite, phenylalanine
production was studied, both in batch culture and with
nongrowing cells of each strain. For this purpose, cells
were transformed with a plasmid encoding the feedback-
resistant enzymes chorismate mutase, prephenate syn-
* Corresponding author. Address: Institute of Biotechnology,
ETH Ho ¨nggerberg, CH-8093 Zu ¨ rich, Switzerland. Telephone: +41/
1/633 3170. Fax: +41/1/633 1051.
420 Biotechnol. Prog. 1998, 14, 420-424
S8756-7938(98)00030-7 CCC: $15.00 © 1998 American Chemical Society and American Institute of Chemical Engineers
Published on Web 05/06/1998