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ý.,,: V. ý ýý . - Nottingham eTheses - University of Nottingham

ý.,,: V. ý ýý . - Nottingham eTheses - University of Nottingham

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Cold adaptation is dependant on the adaptive changes in the cellular proteins and<br />

lipids in response to the conditions imposed on life by such extremely low temperatures<br />

(Russell, 1990). The following sections deal with the definition <strong>of</strong> lower growth-viable<br />

temperature and the range <strong>of</strong> different adaptations to allow psychrophiles to grow at these<br />

temperatures. Mindock et al. (2001) suggest two major physical or physio-chemical<br />

threats to bacterial survival when undergoing freezing:<br />

1. Cell lysis due to increased water volume as ice.<br />

2. Increased salinity outside <strong>of</strong> cell leading to an osmotic gradient across the<br />

membrane.<br />

1.5.1 -<br />

Physio-chemical properties <strong>of</strong> water in confined spaces<br />

Most theories on cold adaptation generally assume that water within a bacterial<br />

cell behaves in the same way as bulk water (Mindock et al., 2001). This however is a<br />

fallacy. Water in confined spaces (e. g. a bacterial cell) has very different properties to<br />

those <strong>of</strong> bulk water systems (Mindock et al., 2001). This is caused by the effect <strong>of</strong> an<br />

ordered surface upon water. The water molecules become highly ordered inducing a<br />

lower freezing point, which alters ionic solubilities, increases viscosity and reduces the<br />

dielectric constants compared to those <strong>of</strong> bulk water (Etzler, 1983). The ordering effect<br />

can extend up to 1 µm from the surface, therefore organising the majority <strong>of</strong> the water<br />

within a cell. The ordering <strong>of</strong> water on a cell membrane can dramatically reduce the<br />

freezing point <strong>of</strong> that water and as such acts as a form <strong>of</strong> cryo-protection. The highly<br />

ordered water near the cell membrane also has reduced solvent properties, thereby<br />

forcing solutes towards the centre <strong>of</strong> the cell causing an increase in solute concentration,<br />

thereby lowering the freezing point <strong>of</strong> the centre <strong>of</strong> the cell (Mindock et al., 2001).<br />

1.5.2 -<br />

Lower growth-viable temperature limit <strong>of</strong> psychrophiles<br />

Reductions in temperature tend to reduce the conformation <strong>of</strong> cellular<br />

macromolecules and other constituents thereby determining the enzymatic reaction rate<br />

(Russell, 1990). This is described in the Arrhenius equation (Herbert, 1986):<br />

K=Ae<br />

-EST<br />

where, Ea is the activation energy, A is a constant relating to steric factors and collision<br />

frequency, R is the universal gas constant, T is absolute temperature (K) and K is the<br />

relation between reaction rate and temperature for a given enzyme.<br />

5

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