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in cyanobacterial FurB/Zur orthologues ( López-Gomollón, Sevilla, Bes,
Peleato, & Fillat, 2009 and Fig. 4.2 ).
3.1.1.2. DNA-binding sites
Under conditions of iron abundance, iron-bound Fur dimers bind to target
promoters in their Fur boxes, also called iron boxes. Fur-binding sequences
were originally regarded as 19-bp inverted repeats with the consensus
(GATAATGATAATCATTATC), and reinterpreted as arrays of NATA/
TAT heptamers ( Escolar, Pérez-Martín, & de Lorenzo, 1999 ). Binding of
FurA from Anabaena to their DNA targets is enhanced in the presence
of divalent metals and reducing conditions ( Hernández, López-Gomollón
et al., 2006 ). Moreover, cyanobacterial FurA binds haeme and this interac-
tion negatively affects its in vitro DNA-binding ability ( Hernández, Peleato,
Fillat, & Bes, 2004 ). Footprinting assays using furA and flavodoxin pro-
moters led to the identification of the first experimentally defined binding
sites for a cyanobacterial FurA protein. These Fur boxes consist of arrays
of A/T-rich sequences that show a faint homology with the Escherichia
coli consensus sequence. Furthermore, experimental work suggests that, in
addition to proper A/T arrays, DNA conformation might be an important
factor in FurA target recognition and binding ( González, Bes, Peleato, &
Fillat, 2011 ). These features are somewhat different from the DNA-binding
properties found for the other cyanobacterial Fur paralogues ( Hernández,
López-Gomollón, Bes, Fillat, & Peleato, 2004 ; López-Gomollón et al., 2009 ;
Napolitano et al., 2012 ).
3.1.2. Occurrence and functions of Fur paralogues in cyanobacteria
The isolation of a fur gene in Synechococcus PCC 7942 through an E. coli -based
in vivo repression assay was the first evidence of the existence of a Fur pro-
tein in cyanobacteria ( Ghassemian & Straus, 1996 ). In this cyanobacterium,
deletion of fur resulted in meridiploids that showed iron-deficiency symp-
toms in iron-replete medium, as well as partial derepression of flavodoxin
and of hydroxamate siderophores. Advances in whole genome sequenc-
ing of diverse cyanobacterial strains allow identification of this essential fur
gene as Synpcc7942_0987. Moreover, searching for ORFs exhibiting the
His-rich motif characteristic of Fur proteins across the 39 cyanobacterial
genomes available in the cyanobase ( http://genome.kazusa.or.jp/cyanobase )
( Nakamura, Kaneko, Hirosawa, Miyajima, & Tabata, 1998 ) reveals the pres-
ence of several Fur paralogues in all the strains sequenced to date. Although
three Fur-like proteins are commonly found in most cyanobacteria, some
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