Mol Gen Genet (1988) 213:487-490
© Springer-Verlag 1988
ISl-mediated mobility of the aerobactin system
of pColV-K30 in Escherichia cob"
Victor de Lorenzo*, Marta Herrero*, and J.B. Neilands
Department of Biochemistry, University of California, Berkeley, CA 94720, USA
Summary. Genes determining the high affinity iron trans-
port system mediated by the siderophore aerobactin are
flanked in the enterobacterial plasmid pColV-K30 by in-
verted repeats of IS/ sequences, suggesting that the aero-
bactin genes are part ofa transposon. To study this possibil-
ity, the entire region between the two IS/ sequences was
cloned as an 18 kb HindIII-BamHI restriction fragment in
pUC8 giving plasmid pMO1. A number of derivatives of
pMO1, in which aerobactin genes were tagged with a kana-
mycin resistance gene, were prepared in order to assess the
ability of both ISis to promote the formation of cointe-
grates with pCJl05, an F derivative devoid of insertion
sequences. Mating-out assays indicated that both flanking
ISis were active in cointegrate formation at detectable fre-
quencies. In some cases, the cointegrates could be resolved,
the final result being a transposition-like event for the entire
aerobactin system.
Key words: pColV - Aerobactin Transposition IS/
Cointegration
Introduction
Aerobactin is an hydroxamate-type siderophore first found
to be produced by Aerobacter aerogenes (Gibson and Ma-
grath 1969). Since the discovery of its production by
Escherichia coli strains carrying the virulence plasmid
pColV-K30 (Warner et al. 1981), systematic surveys in clini-
cal isolates have shown a significant incidence of aerobactin
production in enterobacteria causing different types of in-
fections in humans (Montgomerie et al. 1984; Bindereif and
Neilands 1985; Valvano et al. 1986; Carbonetti et al. 1986).
Aerobactin production has been found in virtually all path-
ogenic enterobacteria (Cercenado etal. 1986; Martinez
et al. 1987).
The genetic organization of the aerobactin determinants
of plasmids pColV-K30 (Carbonetti and Williams 1984;
Roberts et al. 1986a; Ford et al. 1986; de Lorenzo et al.
1986) and pColV-K311 (Gross et al. 1984, 1985)ofE. coli
has been completely ascertained and found to be virtually
identical in both cases. The aerobactin system spans
about 8 kb of DNA and consists of a cluster of five genes
(iucABCD iutA) arrayed in an operon (Fig. 1). The same
genetic arrangement has been detected by DNA hybridiza-
* Present address: Universit6 de Gen6ve, D6partement de Biochi-
mie M~dicale, Centre Medical Universitaire. 9, Avenue de Cham-
pel, CH-1211 Gen6ve 4, Switzerland
Offprint requests to. J.B. Neilands
tion in virtually all aerobactin-producing strains. In addi-
tion to pColV-type plasmids, aerobactin production has
been frequently found to be encoded by non-pColV plas-
mids and, in many cases, is chromosomally determined
(McDougall and Neilands 1984; Marolda et al. 1987; Ro-
berts et al. 1986b).
While the genetic determinants for aerobactin produc-
tion are highly conserved among the different strains which
carry them, the flanking regions have been observed to vary
to different extents. In the prototypical plasmid pColV-
K30, aerobactin genes are flanked by inverted IS1 insertion
sequences and two replication regions (McDougall and Nei-
lands 1984; Perez-Casal and Crosa 1984, 1987). Similar
DNA environments have been found surrounding the aero-
bactin genes of other pColV plasmids, although changes
were observed outside of the approx. 18 kb bracketed by
the insertion sequences and one of the two replication re-
gions was occasionally missing (Waters and Crosa 1986).
In other instances, one of the two insertion sequences and/
or replication regions is missing. Finally, aerobactin genes
can be found devoid of any flanking IS1 or replication
regions (Roberts et al. 1986b; Marolda et al. 1987).
In order to explain the conservation of aerobactin gene
sequences in the midst of variable regions, it has been pro-
posed that in pColV plasmids the region bracketed by ISis
and replication regions could form part of a transposon
(McDougall and Neilands 1984; Perez-Casal and Crosa
1984; Bindereif and Neilands 1985) or of a larger" aerobac-
tin replication unit" (Waters and Crosa 1986). The cases
where these elements are missing (Roberts et al. 1986b;
Marolda et al. 1987), have been claimed to be evidence for
an extinct transposon (Roberts et al. 1986b). However, fail-
ures to show transposition of the system in laboratory-con-
ditions have been reported (Roberts et al. 1986b; Waters
and Crosa 1986).
In this paper, we address the issue of whether IS1 flank-
ing aerobactin genes of pColV-K30 are competent to pro-
mote replicon fusion and, if so, whether that ability may
result in the mobility of the aerobactin system. Genetic evi-
dence shown here indicates that both ISls are indeed able
to induce formation of cointegrates which, upon resolution,
lend a transposition-like character to the aerobactin genes.
Materials and methods
Media, strains and plasmids. Standard LB medium (Miller
1972) was supplemented, when necessary, with 15 gg/ml tet-
racycline (Tc), 60 lag/ml streptomycin (Sin), 50 gg/ml kana-
mycin (Kin), 40 tag/ml choramphenicol (Cm), and 50 ~g/ml
ampicillin (Ap) for monocopy and 200 ~tg/ml for multicopy