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198j 10 Principles, Applications and Future Aspects of Cold-Adapted PGPR<br />

Table 10.1 Percent abundance of membrane fatty acids at different temperatures.<br />

components to remain fluid, a number of changes must occur in the pattern of fatty<br />

acids. Unsaturation of fatty acid chains is the most common change that occurs<br />

when the temperature is reduced; this increases the fluidity of the membrane<br />

because unsaturated fatty acid groups create more disturbance to the membrane<br />

than saturated chains. This process is achieved <strong>by</strong> desaturases situated in the membrane<br />

itself and thus is able to react quickly. In cyanobacteria, four desaturase genes<br />

(desA–desD) have been reported; moreover, desA, desB and desD have been demonstrated<br />

to be cold inducible in Synechocystis [17].<br />

There are, however, a number of other alterations that can occur after a decline in<br />

temperature [18]. The average fatty acid chain length may be shortened, which would<br />

have the effect of increasing the fluidity of the cell membrane because there are<br />

fewer carbon–carbon interactions between the neighboring chains [19]. A psychrophilic<br />

organism, for example Micrococcus cryophilus, which contains high proportions<br />

of unsaturated fatty acids under all growth conditions, responds to a decrease<br />

in temperature, from 20 to 0 C, <strong>by</strong> a reduction in the average chain length of the<br />

fatty acids [20]. All these changes, as summarized in Table 10.1, result in the<br />

membrane maintaining its fluidity <strong>by</strong> producing lipids with a lower gel-to-liquid<br />

crystalline transition temperature and <strong>by</strong> incorporating proportionally more low<br />

melting point fatty acids into membrane lipids.<br />

10.2.2<br />

Carbon Metabolism and Electron Flow<br />

Percent abundance of fatty acids<br />

Fatty acids (IUPAC name) No. of carbon 20 C 18 C 15 C 10 C 5 C<br />

Didecanoic C12 (0) 1.94 2.97 1.18 1.0 0.83<br />

Tridecanoic C13 (0)* 2.08 2.24 2.17 1.46 0.84<br />

n-Tetradecanoic C14 (0) 2.53 3.78 2.57 1.8 0.97<br />

Pentadecanoic C15 (0) 1.50 1.69 1.66 2.41 2.50<br />

cis,cis-9,12-Hexadecanoic C16 (2) 0.65 1.35 1.18 0.67 1.63<br />

cis-9-Hexadecanoic C16 (1) 33.8 33.3 34.7 40.5 49.8<br />

n-Hexadecanoic C16 (0) 27.5 25.4 24.9 20.2 13.2<br />

cis,cis-9,12-Octadecadienoic C18 (2) 0.92 1.08 0.99 2.00 1.00<br />

cis-9-Octadecanoic C18 (1) 21.2 22.5 23.9 26.6 25.9<br />

n-Octadecanoic C18 (0) 7.89 4.30 4.9 3.06 3.31<br />

* Source: Russell [21].<br />

Sardesai and Babu [22] reported that carbon metabolism and electron flow are also<br />

affected <strong>by</strong> low temperature. Cold stress induces a change from respiratory metabolism<br />

to anaerobic lactate formation in a psychrophillic Rhizobium strain. Analysis of<br />

glucose-6-phosphate dehydrogenase and 6-phosphogluconate dehydrogenase of the<br />

pentose phosphate pathway showed an upward regulation of an alternative pathway

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