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normal flora E. coli , EIEC are non-motile and 70% of isolates are unable to
ferment lactose ( Silva et al., 1980 ; van den Beld and Reubsaet, 2012 ). These
are also features of Shigella . More striking is the observation that strains of
EIEC are almost universally negative for lysine decarboxylase (LDC) activity
whereas almost 90% of normal flora E. coli are positive. In this respect, EIEC
also resemble Shigella, which uniformly lack LDC activity. Some serotypes of
EIEC even share identical O-antigens with Shigella ( Sansonetti et al., 1985 ). By
contrast, EIEC resemble E. coli in their ability to ferment xylose and to produce
gas from glucose, both traits for which Shigella are negative ( Silva et al., 1980 ).
It is now generally accepted that the present day strains of Shigella arose
multiple times from as many as seven independent ancestral strains of E. coli
( Pupo et al., 2000 ; Yang et al., 2005 ). EIEC probably evolved later than Shigella
and from different E. coli ancestors ( Lan et al., 2004 ). However, the common
seminal event in the evolution of both groups of pathogens was the acquisition
of a large plasmid that encodes the genes necessary for invasion of mammalian
cells (see below) ( Lan et al., 2001 ).
Evolution of Shigella species and EIEC
The EIEC were recognized as a heterogeneous group of pathogens that resem-
bled Shigella in their pathogenic potential (and in certain metabolic traits) but
were more closely related to E. coli . Early genetic studies demonstrated that
essentially all the virulence factors required for expression of the invasive phe-
notype are encoded on a large plasmid present in all Shigella and EIEC iso-
lates examined ( Sansonetti et al., 1981, 1982b ; Harris et al., 1982 ). Therefore,
from the pathogenesis perspective, EIEC are closely related to Shigella . These
observations led many investigators to propose, incorrectly, that the EIEC rep-
resented a missing link in the evolution from E. coli to Shigella .
Several studies have established that, like the EIEC, the four species of
Shigella are so closely related to E. coli that they should all be included in a
single species. The chromosomes of these organisms are largely co-linear and
are more than 90% homologous ( Brenner et al., 1969 ). Therefore, Shigella , like
EIEC, are a group of pathogenic E. coli. In fact, several investigators using
different approaches have established that the four species of Shigella evolved
from separate E. coli strains ( Pupo et al., 1997, 2000 ; Rolland et al., 1998 ).
Seven different Shigella lineages have been identified through sequence analy-
sis of multiple chromosomal loci. Bacteria expressing the Shigella phenotype
were generated through horizontal transfer of the virulence plasmid from an
unknown donor bacterium to commensal E. coli ( Pupo et al., 2000 ). Thus,
horizontal transfer of the Shigella virulence plasmid to commensal E. coli
has occurred multiple times, each time giving rise to new Shigella clones.
These findings suggest that traits unique to and shared by Shigella species and
EIEC are the result of convergent evolution either through gain-of-function
mutations (e.g. horizontal transfer of the virulence plasmid) or loss-of-function
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