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TF is transiently expressed in mesophilic bacteria but continuously over-
expressed in psychrophiles to achieve cold adaptation, rescuing the chaper-
one function at low temperatures ( Piette et al., 2010, 2011 ).
Either PPiases or TF act as potential CAPs in the proteome of most cold-
adapted microorganisms analysed so far ( Goodchild et al., 2005, 2004;
Kawamoto et al., 2007; Qiu et al., 2006; Suzuki, Haruki, Takano,
Morikawa, & Kanaya, 2004; Ting et al., 2010 ), suggesting that the constraint
imposed by protein folding at low temperature and the cellular responses are
common traits in most psychrophiles ( Piette et al., 2010 ). In contrast, an
almost inverse regulation was found in P. arcticus where GroEL/ES
chaperonins and repression of TF are up-regulated under cold conditions
( Bergholz et al., 2009; Zheng et al., 2007 ). Increased synthesis of
chaperonins has also been reported in S. alaskensis ( Ting et al., 2010 )
possessing two sets of dnaK-dnaJ-grpE gene clusters; proteomic analysis sug-
gests that one of these sets functions as a low-temperature chaperone system
whereas the other functions at higher temperatures ( Ting et al., 2010 ).
At low temperature, in accordance with reduced biomass, almost half of
the down-regulated proteins are involved in general bacterial metabolism.
Most of these proteins are involved in oxidative metabolism, including gly-
colysis, the pentose phosphate pathway, the Kreb's cycle and the electron
chain transporters ( Piette et al., 2011; Wilmes et al., 2011 ).
The Ph TAC125 genome contains genes putatively involved in NO
metabolism, such as NO reductase, PSHAa2417 ,andNO 2 reductase,
PSHAa1477 ( M´digue et al., 2005 ). In this context, the presence of multiple
genes in distinct positions on chromosome I encoding three TrHbs (annotated
as PSHAa0030 , PSHAa0458 , PSHAa2217 )andaFHb PSHAa2880 )
( Giordano et al., 2007; M´digue et al., 2005 ) may be pivotal for cell protection
(see Section 6 ).
5.3. Excess of O 2 and metabolic constraints
Gases (e.g. O 2 ) and radicals (e.g. NO) are highly soluble and stable at
low temperature with visible consequences in genome annotations in
cold-adapted bacteria, having developed responses to strong oxidative stress
(see Casanueva et al., 2010 ).
The apparent benefits of easier O 2 supply are contrasted by the adverse
effects of low temperature on (macro)molecular functions and on the
increased production of RNS and reactive oxygen species (ROS)
( Casanueva et al., 2010; D'Amico et al., 2006 ). In fact, although RNS and
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