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domains ( Pade et al., 2012 ). Corresponding to the occurrence of the otsAB
gene, Crocosphaera accumulates trehalose as only compatible solute. Overex-
pression of this gene in E. coli verified that the gene codes for a functional
trehalose biosynthesis enzyme. Interestingly, C. watsonii WH 8501 acquired
this gene by a lateral gene transfer event ( Pade et al., 2012 ). Cyanobacteria
closely related to Crocosphaera usually accumulate GG, however, these genes
are missing (lost?) in this cyanobacterium.
3.3. Glucosylglycerol
Glucosylglycerol (GG; α-D-glucopyranosyl-(1 → 2)-glycerol) was the first
compatible solute, whose accumulation was found in a cyanobacterium
( Borowitzka et al., 1980 ). Later on, it was detected in many cyanobacte-
rial strains and was regarded to be characteristic for the group of moder-
ate halotolerance ( Hagemann, 2011 ; Reed, Borowitzka et al., 1986 ). While
GG-accumulating beta-cyanobacteria are mostly euryhaline, i.e. they can be
cultivated in freshwater and salt-containing media (e.g. Synechocystis 6803
or Synechococcus sp. PCC 7002; Engelbrecht, Marin, & Hagemann, 1999 ;
Marin, Zuther, Kerstan, Kunert, & Hagemann, 1998 ), the GG-accumulating
picoplanktonic Synechococcus strains, which belong to the alpha-cyanobac-
teria, show only a small range of halotolerance, i.e. they can be cultivated
only in media near the normal seawater salinity level ( Klähn, Steglich et al.,
2010 ). Beside its osmotic function, GG has good direct membrane and
protein stabilizing properties, which are also of biotechnological interest
( Borges, Ramos, Raven, Sharp, & Santos, 2002 ; Hincha & Hagemann, 2004 ;
Sawangwan, Goedl, & Nidetzky, 2010 ).
Cyanobacteria use a two-step biosynthesis for GG ( Hagemann & Erdmann,
1994 ) with an initial GG-phosphate synthase (GgpS) and a subsequent
GG-phosphate phosphatase (GgpP):
(GgpS) ADP-glucose + glycerol 3-phosphate → glucosylglycerol-
phosphate + ADP
(GgpP) Glucosylglycerol-phosphate → glucosylglycerol + Pi
This biosynthetic pathway is similar to that of sucrose, trehalose and glu-
cosylglycerate ( Klähn & Hagemann, 2011 ). However, sucrose and trehalose
biosyntheses prefer UDP-glucose as glucosyl donor, while cyanobacterial
GG synthesis is strictly dependent on ADP-glucose ( Hagemann & Erdmann,
1994 ; Miao et al., 2003 ). Many heterotrophic bacteria also accumulate GG
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