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Accumulation of UV-Absorbing Compounds as a Strategy Against UVR
KUNSHAN GAO AND JUNTIAN XU
They increased in cellular content with increased UV exposure (Brenowitz and
Castenholz, 1997; Pavia et al., 1997; Han and Han, 2005; Zheng and Gao, 2009) to
reduce UV-related photoinhibition and damage, playing a protective role against
solar UVR (Oren and Gunde-Cimerman, 2007).
Seaweeds often exhibit high levels of UVAC, such as MAAs in the red alga
Porphyra columbina (Korbee-Peinado et al., 2004), an unknown UV-B absorbing
substance in the green alga Ulva pertusa (Han and Han, 2005), and phlorotannin
in the brown algae Ascophyllum nodosum and Fucus gardneri (Pavia et al., 1997;
Henry and Van Alstyne, 2004). Higher levels of UVAC have been found in the red
alga Gracilaria lemaneiformis under full spectrum of solar radiation than UVRfree treatments, reflecting a responsive induction (Gao and Xu, 2008). Synthesis
of UVAC has been found to be induced by UV-B in Chondrus crispus (Karsten
et al., 1998), Porphyra columbina (Korbee-Peinado et al., 2004), and Ulva pertusa
(Han and Han, 2005). Such stimulation is dependent on both dose and wavelength, with higher accumulation of UVAC under high daily doses (Karsten et al.,
1998; Franklin et al., 2001). UVR was suggested to trigger some photoreceptors
(active wavelengths between 280 and 320 nm) in the algae to sense the need for
UVAC synthesis (Han and Han, 2005; Oren and Gunde-Cimerman, 2007).
Accumulation of UVAC is often associated with decreased Chl a, resulting in an
increased ratio of UVAC to Chl a (Gao and Xu, 2008).
MAAs, the most common UV-screening compounds, are water-soluble
substances with absorption maxima ranging from 310 to 360 nm (Nakamura
et al., 1982). Although their UVR-protective function is not yet completely clear,
the most acceptable interpretation is that they play a role as a screen against UVR
(Conde et al., 2000; Karsten et al., 2005). Some of these compounds may also
function as antioxidants (Dunlap and yamamoto, 1995; Suh et al., 2003), osmosisregulating substances (Oren, 1997), antenna pigments channeling the energy to the
photosynthetic apparatus (Sivalingam et al., 1976; Gao et al., 2007), or an intracellular
nitrogen storage (Korbee-Peinado et al., 2004; Korbee et al., 2006). Accumulation
of MAAs could be induced by different radiation treatments (Karsten et al., 1999;
Korbee-Peinado et al., 2004; Karsten et al., 2005) or affected by osmotic stress
(Oren, 1997; Klisch et al., 2002) and nutrient availability (Korbee-Peinado et al.,
2004; Korbee et al., 2005; Zheng and Gao, 2009). The accumulation of MAAs was
found to be dependent on both dose and wavelength of incident solar radiation,
with higher accumulation of MAAs associated with high daily doses in Chondrus
crispus (Karsten et al., 1998; Franklin et al., 2001). Nutrient availability was also
found to affect the accumulation of MAAs (Karsten and Wiencke, 1999; KorbeePeinado et al., 2004); enrichment of nitrate enhanced the content of MAAs in
Gracilaria tenuistipitata (Zheng and Gao, 2009). Porphyra plants contain high
levels of MAAs (up to 1% of the dry weight), mainly porphyra-334, which accumulates to the highest concentrations among the species of red algae studied so
far (Gröniger et al., 1999; Hoyer et al., 2001). However, some studies showed that
contents of MAAs did not increase in response to UVR or PAR and could not
ECOLOGICAL AND PHySIOLOGICAL RESPONSES OF MACROALGAE
completely protect Porphyra umbilicalis and Gracilaria cornea against UVR
(Gröniger et al., 1999; Sinha et al., 2000).
Distribution of seaweed at different zonational depths affects the accumulation of MAAs. Intertidal species are usually more resistant to UV stress (i.e.,
inhibition of photosynthesis) than subtidal species that have less or no MAAs
(Maegawa et al., 1993). It was found that deep-water polar macroalgal species did
not have MAAs, whereas supra- and eulittoral species contained MAAs to high
concentrations (Hoyer et al., 2002). Total MAAs content in Mastocarpus stellatus
was sixfold higher than in Chondrus crispus that was generally found at a greater
depth; quantum yield and maximal electron transport rate were more reduced in
C. crispus than M. stellatus by UV-B radiation (Bischof et al., 2000). MAAs content in Devaleraea ramentacea increased with decreased depth, being correlated
with a higher photosynthetic capacity under UVR treatment (Karsten et al.,
1999). The macroalgal zonation patterns relate to their ability to resist high
radiation stress (Hanelt, 1998).
Macroalgal species distributed at the upper part of intertidal zone may be
exposed to much higher solar radiation during emersion if the low tide coincides
with local noon. Recently, it was shown that desiccation or dehydration of
Porphyra haitanensis thalli led to higher absorptivity of the UVAC (Jiang et al.,
2008). The ability for Porphyra haitanensis thalli to increase its cellular content of
UVAC during such emersion period allows them to cope with UVR stress. The
possible strategy for macroalgal species to survive at the upper levels of intertidal
zone is to increase its content of UVAC, which play roles in both sunscreening
and osmosis regulation.
PAR drives photosynthesis, whereas UVR is usually known to harm physiological
processes in macroalgae as well as phytoplankton. UV-A, however, at reduced
levels, has been shown to enhance photosynthesis and repairing processes of photodamaged molecules, whereas UV-B mostly results in harmful effects. During their
long history of evolution, seaweeds have developed protective strategies against
harmful UV irradiances, such as synthesizing and accumulating UVAC and the
repair of DNA damage. Different life stages of seaweeds show different sensitivity
to solar UVR, with less-differentiated forms being more sensitive to UVR. Species
distributed at different depths in the intertidal zone also show different responses
to solar UVR; upper species, that are usually exposed to higher levels of solar
radiation and accumulate higher contents of UVAC (such as MAAs), are more
tolerant of UVR. On the other hand, diurnal photosynthesis can be underestimated during twilight period or cloudy days and overestimated during noontime
if the effects of UVR are ignored owing to positive and negative effects caused by
UV-A, respectively, at low and high irradiance levels.
KUNSHAN GAO AND JUNTIAN XU
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Biodata of E. Walter Helbling, Virginia E. Villafañe, and Donat-P. Häder,
authors of “Ultraviolet Radiation Effects on Macroalgae from Patagonia, Argentina”
Dr. E. Walter Helbling is currently the Director of Estación de Fotobiología
Playa Unión (EFPU, Argentina) and a researcher from Consejo Nacional de
Investigaciones Científicas y Técnicas (CONICET, Argentina). He obtained his
Ph.D. from Scripps Institution of Oceanography, University of California, San
Diego (USA). His scientific interests are in ecophysiology of plankton and photobiology of aquatic systems in relation to climate change.
Dr. Virginia E. Villafe is currently a Researcher from Consejo Nacional de
Investigaciones Científicas y Técnicas (CONICET, Argentina). She obtained
her Ph.D. from University of Groningen (The Netherlands) and continued her
research in Patagonia at Estación de Fotobiología Playa Unión (EFPU, Argentina).
Dr. Villafe´s scientific interests are in the areas of ecophysiology of plankton and
Virginia E. Villafañe
E. Walter Helbling
A. Israel et al. (eds.), Seaweeds and their Role in Globally Changing Environments,
Cellular Origin, Life in Extreme Habitats and Astrobiology 15, 199–214
DOI 10.1007/978-90-481-8569-6_12, © Springer Science+Business Media B.V. 2010
E. WALTER HELBLING ET AL.
Professor Donat-P. Häder holds the Chair of Plant Ecophysiology at the Department
for Biology at the Friedrich-Alexander University in Erlangen-Nürnberg. He
obtained his Ph.D. from the University of Marburg in 1973. After a Postdoc year
in East Lansing Michigan state, he became Researcher in Marburg. Professor
Häder’s scientific interests are in the areas of the effects of stratospheric ozone
depletion and resulting increasing solar UV-B radiation at the Earth’s surface on
the biota. He concentrates on these effects in combination with global climate
change on aquatic ecosystems in many habitats over the globe.