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environment at their natural growth site. Furthermore, the accumulation of DNA
damage in seaweeds can be regarded as a very good measure for the degree of
adaptation of a given species towards UVB exposure (van de Poll et al. 2001 ).
These findings were also confirmed in a transplantation experiment on another
deep-water red algal species Delesseria sanguinea , in which growth was also
impaired when the alga was exposed to surface solar radiation (Pang et al. 2001 ).
Results obtained indicated the absence of any protecting mechanisms against
excessive radiation in this algal species, which are, however, not relevant to the
species as it inhabits the low light environments in the shade of canopy algae or in
great water depths. Thus, sublittoral seaweeds may avoid being exposed to UVB
simply by growing in water depths, in which most of the UVB irradiance is already
attenuated by the water column above. A prominent example of this avoidance
strategy is also the Arctic endemic deep-water kelp Laminaria solidungula (Brey
2009 ). Other algal species following this strategy may also be growing in the
understory being protected by the algae above or following a cryptic life cycle.
Generally, all modulation of ecophysiological reactions toward variation in
abiotic factors is conditioned by genetic adaptation. This is also visible in two red
algal species from Spitsbergen (Arctic) with slightly different vertical zonation
preferences (Karsten et al. 1999 ): Devaleraea ramentacea as a species from
shallow waters is permanently equipped with high activities of reactive oxygen
scavenging superoxide dismutase (SOD, see Chap. 6 by Bischof and Rautenberger).
This high but static activity is reasonable for a species from shallow waters, where
usually strong variation in abiotic conditions and, thus, the onset of stressful
conditions to photosynthesis potentially resulting in increased ROS production is
more likely than in more stable deeper waters. However, maintaining a protective
system on such a high level throughout the year is probably energetically cost
intense. Thus, species which are not permanently exposed to stressful conditions,
e.g., in deeper waters, may favor the strategy to rather respond to abiotic stress and
to increase protective strategies, like SOD activity, only when they are needed
during times of, e.g., high UV irradiance. Palmaria palmata inhabiting slightly
deeper waters than D. ramentacea is applying this strategy (Karsten et al. 1999 ;
see also Bischof et al. 2006a ).
20.3 Acclimation to UV Exposure in Space and Time: Depth
Distribution, Small-Scale Gradients, and Season
Seaweeds populating a flexible environment, i.e., the shallow water zones,
coastlines with strong seasonality, etc., require mechanisms of acclimation in
order to set physiological performance to the variation of environmental
requirements. Thus, also acclimation toward UV exposure has been widely
observed in seaweeds. Acclimation of photosynthesis to UV exposure has been
demonstrated in the Arctic/cold-temperate kelp Alaria esculenta from Spitsbergen
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