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under saline conditions. These organisms use ADP- and also UDP-glucose
for the GgpS reaction. Moreover, in many heterotrophic bacteria, the two
enzyme activities, GgpS and GgpP, are found in one continuous protein
( Hagemann et al., 2008 ). Such fused GG synthesis proteins have not yet
been found in any cyanobacterial genome.
Genome searches using the GgpS protein (Sll1566) from Synechocys-
tis 6803 ( Marin et al., 1998 ) identified 28 genes coding for highly similar
proteins in other cyanobacteria ( Table 2.1 ). The closest homologue is the
GgpS from Synechococcus sp. PCC 7002, which has been functionally veri-
fied ( Engelbrecht et al., 1999 ). Among beta-cyanobacteria, it is also found
in many euryhaline and marine strains such as Arthrospira ( Yoshikawa et al.,
2011 ), Acaryochloris , and some but not all Cyanothece strains ( Table 2.1 ).
However, there are also many marine strains, which do not harbour ggpS
genes in their genomes and are accordingly not using GG as main compat-
ible solute. Interestingly, the marine mat-forming Microcoleus chtonoplastes
contains genes for GG synthesis, whereas the terrestrial M. vaginatus strain
FGP-2 is only able to synthesize trehalose ( Table 2.1 ). Probably, in the ter-
restrial habitat, desiccation is the main stress, which is usually tolerated by
trehalose accumulation, while the marine Microcoleus is faced by salt stress,
therefore preferring GG.
As previously reported ( Scanlan et al., 2009 ), practically all of the marine
picoplanktonic Synechococcus strains (only exception is Synechococcus sp. WH
7805) contain ggpS genes, whereas the related Prochlorococcus strains miss
it all ( Table 2.1 ). The accumulation of GG in marine Synechococcus and its
absence in Prochlorococcus strains have been recently verified ( Klähn, Steglich
et al., 2010 ).
The GgpP (StpA was used synonymously) protein is a specific GG-
phosphate phosphatase. Homologues of this protein are restricted to
cyanobacteria and are characterized by a specific protein domain called
Salt_tol_Pase superfamily. The only functional characterized GgpP
enzyme is encoded by slr 0746 in Synechocystis 6803 ( Hagemann, Schoor,
Jeanjean, Zuther, & Joset, 1997 ). Highly similar proteins are found in all
cyanobacterial genomes harbouring a ggpS gene. Among beta-cyanobac-
teria ggpS and ggpP never form an operon. However, in the oceanic pico-
planktonic Synechococcus strains, these genes coding for functional linked
proteins are often found adjacent to each other indicative for operon
structures.
In one case, Synechococcus sp. RCC307, not only ggpS and ggpP are linked
but also the four subunits for the putative ABC-type GG-transport system
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